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Full text of "Rhodora [electronic resource]"

JOURNAL OF THE 



NEW ENGLAND BOTANICAL CLUB 




Vol. 77 



March, 1975 



No. 809 



®lfe 'pizfo ^nglattii potato*! QHnlt, <3Ittc 

Botanical Museum, Oxford Street, Cambridge, Mass. 02138 



Conducted and published for the Club, by 
ALFRED LINN BOGLE, Editor-in-Chief 

ROLLA MILTON TRYON 
STEPHEN ALAN SPONGBERG 
GERALD JOSEPH GASTONY 
RICHARD EDWIN WEAVER 



Associate Editors 



RHODORA. — A quarterly journal of botany, devoted primarily to 
the flora of North America and floristically related areas. Price 
$20.00 per year, net, postpaid, in funds payable at par in the United 
States currency at Boston. Some back volumes, and single copies 
are available. For information and prices write RHODORA at 
address given below. 

Scientific papers and notes, relating directly or indirectly to the 
plants of North America, will be considered by the editorial com- 
mittee for publication. Articles concerned with systematic botany 
and cytotaxonomy in their broader implications are equally accept- 
able. All manuscripts should be submitted in duplicate, and should 
be double-spaced throughout. Please conform to the style of recent 



issues of the journal. Illustrations can be used only if the cost of 
engraver's blocks is met through the author or his institution. 
Forms may be closed five weeks in advance of publication. Ex- 
tracted reprints, if ordered in advance, will be furnished at cost. 

Address manuscripts and proofs to: 

Dr. A. Linn Bogle 

Dept. of Botany and Plant Pathology, 
University of New Hampshire, 
Durham, New Hampshire 03824 

Subscriptions and orders for back issues (making all remittances 
payable to RHODORA) should be sent to RHODORA, Botanical 
Museum, Oxford Street, Cambridge, Mass. 02138. In order to receive 
the next number of RHODORA, changes of address must be received 
prior to the first day of March, June, September or December. 

Second Class Postage Paid at Boston, Mass. 



MANUFACTURED BY 

THE LEXINGTON PRESS. TNC. 
LEXINGTON, MASSACHUSETTS 



IRbofcora 



JOURNAL OF THE 



NEW ENGLAND BOTANICAL CLUB 



Vol 77 March, 1975 No. 809 



ON THE EPIBIOTIC AND PELAGIC 

CHLOROPHYCEAE, PHAEOPHYCEAE, 

AND RHODOPHYCEAE OF THE 

WESTERN SARGASSO SEA 



William J. Woelkerling 



This paper provides a taxonomic account of the epibiotic 
and pelagic Chlorophyceae, Phaeophyceae, and Rhodophy- 
ceae collected during six cruises to the Western Sargasso 
Sea and follows two previous studies by the author (Woel- 
kerling, 1972; 1973) of non-planktonic algae from this re- 
gion. Earlier published records (e.g. Collins, 1917; Conover 
& Sieburth, 1964; Farlow, 1914; Hentschell, 1921; Pratt, 
1935; Winge, 1923) of green and red algae and of brown 
algae other than Sargassum from the Sargasso Sea are 
few and fragmentary, and, with one. or two exceptions, they 
do not include identifications to species level. 

Nearly 75 percent of the taxa encountered during this 
investigation have not been reported previously from the 
Sargasso Sea, and these new records raise the total known 
flora of the region to include 10 Chlorophyceae, 25 Phaeo- 
phyceae, and 33 Rhodophyceae. Epibiotic Bacillariophyceae 
(see Carpenter, 1970) and epibiotic Cyanophyceae (see 
Carpenter, 1972; Hentschell, 1921) are not treated in this 
paper. 



1 



2 



Rhodora [Vol. 77 



Methods of sampling and processing are outlined else- 



where (Woelke 



(with speci- 



men numbers prefaced by WJW) has been retained in the 
author's personal collections, currently housed at wis. 
Other herbarium abbreviations follow Lanjouw & Stafleu 
(1964). 

Data provided for each taxon includes references to 
records of occurrence in adjacent regions and/or of general 
taxonomic value, the type locality and reported location of 
the type collection (in most cases, location of types has not 
been verified), known distribution based on published rec- 
ords, and collection data for all specimens examined. This 
information usually is followed by ecological and/or taxo- 
nomic notes. In cases where specific identification has not 
been possible due to fragmentary and /or very young or 
small plants, the available data has been summarized briefly 
at the generic level. The genus Sargassum presents special 
problems regarding species identification, and these are 
outlined in the discussion of that taxon. 

Epibiotic taxa can be divided conveniently into two eco- 
logical groups, namely the permanent element and the in- 
vading element. The former includes all taxa epibiotic on 
Sargassum fluitans and/or S. nutans, the two brown algae 
which comprise the vast bulk of Sargasso Sea vegetation, 
estimated by Parr (1939) to be in excess of 4 X 10 r! metric 
tons. These two species apparently are endemic to the 
Sargasso Sea. The invading element includes all other 
"macroscopic" taxa (Ascophyllum, Fucks and their asso- 
ciated epiphytes (Woelkerling, 1972), species of Sargassum 
other than S. fluitans and S. natans, etc.). 

The permanent epibiotic element comprises both taxa 
which are normally of small size (less than 1 cm tall) and 
diminutive forms of taxa normally of larger stature. Spe- 
cies in both groups, however, frequently bear reproductive 
structures, in marked contrast to Sargassum fluitans and 
5. natans which never have been found with receptacles 
(see, however, Parr, 1939, page 49) and reproduce solely 
by fragmentation as far as is known. 



1975] Sargasso Sea — Woelkerling 3 



Sincere thanks are due Mr. Gordon Volkmann of the 
Woods Hole Oceanographic Institution for making arrange- 
ments for the collection of samples in the Sargasso Sea and 



for mak 
cruises. 



M 



am 



DIVISION CHLOROPHYTA 

Class Chlorophyceae 

order tetrasporales 

1 

Family Palmellaceae 
Genus Pseudotetraspora Wille, 1906 
Pseudotetraspora marina Wille 1906:20, Taf. 1, Figs. 32-36. 
TYPE locality: Steinviksholm, Drontheimsfjord, Norway. 

TYPE : not located. 

DISTRIBUTION: apparently known only from the Sargasso 
Sea and Norway. 

specimens examined: Sargasso Sea: 31°N-70°W, 5.vii.l970, Volk- 
ma'nn. (WJW 2735); 32°09'N-(;4°58'W, lf>.v.l<>70, Woclkerlmg (WJW 
2G70); 34°N-70 1 W, 30.vi.1970, Volkmartm (WJW 2727), r>.vii.l970, 
Volkmann (WJW 2748); 35°54'N-70°30'W, 13.viii.1970, Moore (WJW 
2901); 37°N-70°W. 12.V.1970. Woelkerling (WJW 2(525); 37°30'N- 
70°W, 8.vii.l970. Volkmann (WJW 2708); 39°07'N-70°35'W, IG.viii. 
1970,'j/oo,v (WJW 2919, 2927); 39°30'N-71°W, fi.x.1970, Volkmann 



(WJW 2871). 



am 



as epiphytes on Sargassum nutans, Sargassum sp., and 

Cladophora. 

Howe (1920) described a Pseudotetraspora antilJxirum 

from the Bahamas and separated it from P. marina on the 
basis of apparent differences in color, shape of the gela- 
tinous mass, and cell size. These criteria require further 
consideration since the Sargasso Sea specimens could con- 
veniently be placed in either taxon. Howe (1920), for ex- 



met 



Wille (1906) 



na 



The Sargasso Sea plants have cell diameters ot 3-14 /xm 



comparison 



them 



4 



Rhodora [Vol. 77 



ORDER CHAETOPHORALES 

Family Chaetopeltidaceae 

Genus Diplochaete Collins, 1901 

Diplochaete solitaria Collins 1901:242. Chapman 1961:69, 
Fig. 73. Collins 1909:278, Fig. 99. Collins and Hervey 
1917:38. Taylor 1960:53. 
type locality : Kingston, Jamaica. 

type : NY. 

distribution : Bermuda, Jamaica, Sargasso Sea. 

specimens examined: Sargasso Sea: 26°57'N-72°58'W, 26.iv.1970, 
Moore (WJW 2647, 2651); 28°N-70°W, 4.iii.l970, Volkm'ann (WJW 
243.")); 31°N-70°W, 5.vii.l970, Volkmann (WJW 2734); 32°09'N-64° 
58'W, 16.V.1970, WoeVcerlvng (WJW 2664, 2668) ; 34°N-70°W, 30.vi. 
1970, Volkmann (WJW 2726, 2747); 3r>°r>4'N-70°30'W, 13.viii.1970, 
Moore (WJW 2900); 36°28'N-70°29'W, 15.viii.1970, Moore (WJW 
2908); 37°N-70°W, 12.V.1970, WoelkerUng (WJW 2618); 38°22'N- 
70°58'W, 12.X.1970, Volkmann (WJW 2891); 39°07'N-7o'°35'W, 16. 
viii.1970, Moore (WJW 2916); 39°30'N-71°W, 6.X.1970, Volkmann 
(WJW 2860, 2885). 



common com 



of the Sargasso Sea flora and has been found, sometimes in 
considerable numbers, on both Sargassum fiuitans and S. 
natans as well as on a variety of red algae, Cladophora 
(Chlorophyta), and on hydroids. 

Cells in the collections examined bear 1-4 setae that may 

be oriented in any direction relative to one another but 

generally are directed away from the substrate. As noted 

by Collins (1909) the freshwater taxa often referred to 

the genus Polychaetophota West and West (1903) may be 

congeneric with Diplochaete (see also Printz, 1964; G. S. 
West, 1908). 



ORDER ULVALES 

Family Ulvaceae 

Genus Entcromorpha Link, 1820 
Enteromorpha sp. 

Two collections [31°N-69°29'W, 3.iii.l970, Volkmann 
(WJW 2369) and 34°N-70°W, 7.iii.l970, Volkmann (wjw 
2397)] of Sargassum fiuitans contained epiphytic plants 



1975] Sargasso Sea — Woelkerling 5 



of Enter vmorpha which appear similar to E. flexuosa (Wul- 
fen ex Roth) J. Agardh (see Bliding, 1963, for a detailed 
account of E. flexuosa). Since the Sargasso Sea plants were 
all stunted (under 3 cm tall), however, specific determina- 
tion could not be made with certainty. 

Genus Mono stroma Thuret, 1854 

Monostroma pulchrum Farlow 1881:41. Collins 1909:211. 
Collins, Holden, and Setchell 1900:658. Taylor 1957:72. 

type locality : Watch Hill, Connecticut. 

type: fh. 

distribution : Connecticut to Nova Scotia, Sargasso Sea. 

specimens examined: Sargasso Sea: 39°05'N-69°48'W, lO.v.1970, 
Woelkerling (WJW 2572); 39°11'N-69°24'W, lO.v.1970, Woelkerling 
(WJW 2550). 

Two small plants of Monostroma pulchrum were found 
as epiphytes on Fucus vesiculosus L., which had apparently 
drifted out into the Northwestern fringes of the Sargasso 
Sea (see Woelkerling, 1972). Critical studies are needed 
to determine whether M. pulchrum is really specifically 
distinct from the more widely distributed and better known 
M. oxyspermum (Kuetzing) Doty (see Bliding, 1968, p. 585, 
under Ulvaria oxysperma) . 

Genus Per curs aria Bory, 1823 

Percursaria percursa (C. Agardh) Rosenvinge 1893:963. 
Bliding 1963:20, Figs. 5-6. Collins 1909:197. Kylin 
1949:16, Fig. 9. Taylor 1957:61; 1960:54. 

Enter vmorpha percursa (C. Agardh) J. Agardh Chap- 
man 1961:66, Fig. 70. 
type locality : Adriatic Sea. 

TYPE: LD. 

distribution : widely distributed. 

specimens examined: Sargasso Sea: 34°N-70°W, 13.V.1970, Woel- 
kerling (WJW 2612); 37°N-70°W, 12.V.1970, Woelkerling (WJW 
2627). 

In both cases the host plant was Sargassum natans. The 
main axes and laterals were pleuriseriate while the smaller 
laterals were uniseriate in the specimens examined. 



6 



Rhodora [Vol. 77 



Genus Ulva Linneaus, 1753 



Ulva sp. 

One young plant [34°N-70°W, 7.iii.l970, Volkmann 
(wjw 2412)] of Ulva was found as an epiphyte on a frag- 
ment of the grass Spartina. Specific determination was not 
possible. The Spartina fragment apparently had drifted 
out from the U. S. coast. 



ORDER CLADOPHORALES 

Family Cladophoraceae 

Genus Cladophora Kuetzing, 1843 

As noted by Taylor (1960, p. 78), the genus Cladophora 
has been difficult to cope with in the American tropics and 
critical studies are badly needed. The recent monographs 
of Soderstrom (1963) and van den Hoek (1963), which 
will probably provide a basis for such studies, have been 
used in making specific determinations during this investi- 
gation. 

Cladophora dalmatica Kuetzing 1843:263. van den Hoek 
1963:186, Figs. 601-35. 

C. oblitterata Soderstrom 1963:47, Figs. 38-54A. 

type locality: Split (Spalato), Yugoslavia. 
type: l (No. 937/281/406). 
distribution : probably widespread. 

specimens examined: Sargasso Sea: 26°57'N-72°. r >8'W, 2fi.iv.1970, 
Moore (WJW 2fi4fi) ; 31°N-69°29'W, 3.iii.l970, Volkmann (WJW 

2874); 34°N-70°W, 10.L1970, Volkmann (WJW 2205), 13.V.1970, 
WoeUcerling (WJW 2614) ; 37°N-70°W, 12.V.1970, Woelkerling (WJW 
2fi20); 37°30'N-70°W, 8.vii.l970, Voibmann (WJW 2704); 38°22'N- 
70°58'W, 12.x-.1970, Volkmann (WJW 288fi); 39°07'N-70°3r)'W, 16. 
viii.1970, Moore (WJW 2918). 



mens un to 3 cm 



fj as sum ft 



In all cases the apical cells 



were under 20 f im in diameter and the branch systems 
showed a distinct acropetal organization. 

Cladophora laetevirens (Dillwyn) Kuetzing 1843:267. van 
den Hoek 1963:128, Figs. 409-429, 433, 440. 



1975 ] Sargasso Sea — Woelkerling 7 



type locality: England. 

neotype: bm (H4351/60/6) ; see van den Hoek 1963; 
p. 128. 

distribution : probably widespread. 

specimen examined: Sargasso Sea: 31°N-69°29'W, 3.iii.l970, 
Volkmartn (WJW 2359). 



1 he only specimen (about 2.5 cm tall) encountered during 
this study occurred as an epiphyte on Sargassum fluitans. 

Cladophora socialis Kuetzing 1849:416, 1854:15, pi. 71, 
Fig. 1. van den Hoek 1963:43, Figs. 79-91. 

type locality: Tahiti. 

type: l (937/253/440). 

distribution: Europe, Tropical Oceania, Sargasso Sea. 

specimen examined: Sargasso Sea: 26°57'N-72°58'W, 26.iv.1970, 
Moore (WJW 2650). 



cm 



gaasswn natans. 



Genus Spongomorpha Kuetzing, 1843 

Spongomorpha arcta (Dillwyn) Kuetzing 1849:417. Collins 
1909:359. Taylor 1957:90, pi. 6, Figs. 5-6. 

Cladophora arcta (Dillwyn) Kuetzing 1843:263. Collins, 
Holdenand Setchell 1896:224; 1901:815. 

TYPE locality: England. 
TYPE : presumably in nmw. 

distribution : colder waters of Europe and North Amer- 
ica. 

specimens examined: Sargasso Sea: 38°53'N-69°39'W, lO.v.1970, 
Woelkerling (WJW 2561, 2567); 39 05'N-69°48'W, lO.v.1970, Woel- 
kerling (WJW 2571). 



imen 



of Ascophyllum nodosum and Fucus vesiculosus which had 
drifted out to the Northwestern fringes of the Sargasso 
Sea (see Woelkerling 1972). Spongomorpha arcta is prob- 
ably not a permanent component of the Sargasso Sea flora. 



8 Rhodora [Vol. 77 



DIVISION CHROMOPHYTA 

Class Phaeophyceae 

order ectocarpales 

Family Ectocarpaceae 

Genus Ectocarpus Lyngbye, 1819 

Ectocarpus elachistaeformis Heydrich 1892:470, pi. XXV, 
Fig. 14. Boergesen 1914:18, Fig. 11; 1920:435. Collins 
and Kervey 1917:70. Earle 1969:132, Fig. 28. Taylor 
1928:107, pi. 14, Fig. 12; 1960:202, pi. 29, Fig. 9. 

type locality: New Guinea. 



1 



YPE : probably destroyed. 



distribution: Caribbean Islands, Gulf of Mexico, New 
Guinea, Sargasso Sea. 

specimens examined: Sargasso Sea: 31°N-69°29'W, 3.iii.l970, 
Volkmann (WJW 2382, 2384); 34°N-70°W, 10.i.l970, Volkmann 
(WJW 2211). 



mm 



sum 



gas sum sp. Pluri- 



as 



gate-lanceolate as described by Boergesen (1914) . 



Genus Feldmannia, Hamel, 1939 

Feldmannia irregularis (Kuetzing) Hamel 1931-1939 :XVII, 
Fig. 61F. Cardinal 1964:54, Fig. 29. Kuckuck 1963:371, 
Fig. 6. 

Ectocarpus irregularis Kuetzing. Boergesen 1926:25, 
Figs. 12-14. Chapman 1963:11. Rosenvinge et Lund 
1941 :50, Figs. 23-24. Sauvageau 1933:101, Figs. 24-27. 

type locality : Adriatic Sea. 

type: L. 

distribution: northern Europe, Canary Islands, Sar- 
gasso Sea. 

specimens examined: Sargasso Sea: 28°N-70°W, 4.iii.l970, Volk- 
mann (WJW 2431); 31°N-60°29'W, 3.iii.l970, Volkmann (WJW 
2366); 34°N-70°W, 7.iii.l970, Volkmann (WJW 2413). 



1975] Sargasso Sea — Woelkerling 9 



The Sargasso Sea specimens occur as epiphytes on Sar- 
gassum fluitans and S. natans and agree well with the ac- 
count of Feldmannia irregularis given by Cardinal (1964). 
Plurilocular sporangia are common. Chapman (1963) re- 
cords this taxon (as Ectocarpus irregularis) from Jamaica 
and, following Boergesen (1941), regards E. rallsiae ( 
Giffordia rallsiae (Vickers) Taylor (1960, p. 208), a widely 
distributed taxon in the American tropics) as conspecific. 
Earle (1969), however, maintains G. rallsiae as a distinct 
species. The precise relationships of G. rallsiae and Feld- 
mannia irregularis thus remain uncertain and further criti- 
cal studies of the type and other collections of the two taxa 
are needed. 

Genus Giffordia Batters, 1893 
Giffordia conifera (Boergesen) Taylor 1960:207. Earle 



Col- 



1969:135, Fig. 21. 

Ectocarpus coniferus Boergesen 1914:8, Figs. 5, 6. 

lins and Hervey 1917 :69. 
type locality : U. S. Virgin Islands. 

type: c. 

distribution: Sargasso Sea and tropical western Atlan- 
tic. 

specimens examined: Sargasso Sea: 28°N-70°W, 4.iii.l970, Volk- 
mann (WJW 2430); 34°N-70°W, 7.iii.l970, Volkmann (WJW 2416). 

The Sargasso Sea specimens occur epiphytically on Sar- 
gassum fluitans and S. nutans and agree with the original 
account of Boergesen (1914). Only plurilocular sporangia 
are present. Hamel (1931-39) considers this taxon to be 



maintains 



with Feldmannia irregularis, but Earle (1969) 
it as a distinct species. The status of the taxon 
will remain questionable until a critical study of all the 
ectocarpoid algae of the western tropical Atlantic is un- 
dertaken. 

Giffordia mitchelliae (Harvey) Hamel 1939 :XIV, Fig. 61c, d. 
Cardinal 1964:45, Fig. 23. Earle 1969:138, Fig. 24. 
Taylor 1960:206, pi. 29. Figs. 1-2. 



10 Rhodora [Vol. 77 



Ectocarpus mitchelliae Harvey 1852, p. 142, pi. 12 g. 

Boergesen 1914:6, Figs. 3-4. 1941 :7, Figs. 1-5. Collins 
andHervey 1917:69. 

type locality: Nantucket Island, Massachusetts. 

type: TCD. 

distribution : widespread in tropical and temperate seas. 

specimens EXAMINED: Sargasso Sea: 31°N-69°29'W, 3.iii.l970, 
Votkmann (WJW 2361, 2381, 2383); 32°09'N-64°58'W, 16.V.1970, 
Woelkerling (WJW 2666); 34°N-70 C W, lO.i.1970, Volkma'nn (WJW 
2206, 2218); 39°07'N-70°35'W, 16.viii.1970, Moore (WJW 2925). 

Specimens up to 5 cm tall occur as epiphytes on Sargas- 
sum fluitans, S. natatis, and on an unidentified species of 

Sargassum. In all cases plurilocular sporangia are abun- 
dant. 

Giffordia sandriana (Zanardini in Kuetzing) Hamel 1939: 
XIV. Cardinal 1964:37, Fig. 18. Kylin 1947:10, Fig. 3. 
Taylor 1960 :207. 

Ectocarpus sandrianus Zanardini in Kuetzing 1849:451. 
Rosenvinge et Lund 1941 :44, Fig. 18. 

type locality : Adriatic Sea. 
type: L. 

DISTRIBUTION : Bermuda, Europe, Sargasso Sea. 

specimens examined: Sargasso Sea: 34°N-70°W, 7.iii.l970, Volk- 
mann (WJW 2414). 

Plants up to 5 cm tall occurred as epiphytes on Sargas- 



fl 



Genus Pylaiella Bory, 1823 



Pylaiella littoralis (L.) Kjellman. Cardinal 1964:11, Fig. 1. 
Rosenvinge et Lund 1941:5. Taylor 1957:102, pi. 9, Figs. 
1-3. Woelkerling 1972:298. 

type locality: Europe. 

type: linn. 

distribution : widespread. 

specimens examined: Sargasso Sea: 38°53'N-69°39'W, lO.v.1970, 
Woelkerling (WJW 2559); 39°11'N-69°24'W, lO.v.1970, Woelkerling 
(WJW 2555). 



1975] Sargasso Sea — Woelkerling 11 



The plants were found as epiphytes on Fucus vesiculosus 
along the northwest fringes of the Sargasso Sea and prob- 
ably should not be considered as a permanent component of 
the Sargasso Sea flora (Woelkerling, 1972). 



ORDER SPHACELARIALES 

Family Sphacelariaceae 

Genus Sphacclaria Lyngbye, 1819 

Sphacelaria fucigera Kuetzing. Sauvageau 1901:145, Fig. 
35. Taylor 1960:210, pi. 29, Fig. 5. Womersley 1967:199, 

TYPE locality: Karak Island, Persian Gulf. 

type: l (937/71/472). 

distribution: cosmopolitan in tropical and temperate 
waters. 

specimens examined: Sargasso Sea: 34°N-70°W, lO.i.1970, Volk- 
mann (WJW 2202); 38°22'N-70°58'W, 12.X.1970, Volkmann (WJW 
2896); 39°07'N-70°35'W, 16.viii.1970, Moore (WJW 2921). 

The plants were epiphytic on Sargassum natans and an 
unidentified Sargassum and bore numerous propagula. One 
collection (WJW 2921) also had numerous multicellular 



hairs. 



ORDER DICTYOTALES 

Family Dictyotaceae 
Genus Dictyota Lamouroux, 1809 



Dictyota sp. 



Two collections [32°09'N-64°58'W, 16.V.1970, Woelh 
ling (WJW 2658) and 39°30'N-71°W, 6.X.1970, Volknu 
(WJW 2877)] of small plants referrable to Dictyota h, 
been made during this study. The former, which contaii 
both male and female reproductive structures and was ab 
5 cm tall, was found growing on a fragment of Colpomei 
which nrobablv became detached and drifted out from 



mu 



umber 



dichotoma. but definite specific determination 



was not considered possible. 



12 Rhodora [Vol. 77 



The latter plants were attached to a fertile piece of Sar- 
g as sum (origin uncertain) and were very young (less than 
2 cm tall) ; specific determination likewise could not be 
made. It seems more probable that Dictyota is an invader 
rather than a permanent component of the Sargasso Sea 
flora considering that both host plants themselves appear 
to be invaders. 

Prat (1935, p. 128) makes mention of a Dictyota cervi- 
cornis from this region, but no specimens referrable to that 
species have been encountered during the present study. 



Genus Padina Adanson, 1763 



Padina sp. 



>8'W, 

piec^ 



Woelkerling (WJW 2657)] occurred on a 
menia which apparently had drifted out from Bermuda. 
Specific determination was not possible and it seems prob- 
able that Padina is an invader rather than a permanent 
component of the Sargasso Sea flora. 



ORDER CHORDARIALES 

Family Elachisteaceae 

Genus Elachistea Duby, 1830 

Elachistea lubrica Ruprecht. Collins, Holden, and Setchell 
1898:480. Taylor 1957:140. Woelkerling 1972:297. 

type locality : Okhotsk Sea. 

type: le. 

distribution: reported from eastern North America, 
Greenland, and the Okhotsk Sea. 

specimens examined: Sargasso Sea: 38°53'N-69°39'W, lO.v.1970, 
Woelkerling (WJW 2564); 39°05'N-69°48'W, lO.v.1970, Woelkerling 
(WJW 2569); 39°11'N-69°24'W, lO.v.1970, Woelkerling (WJW 2547; 
2556). 

An invader species attached to drifting Ascophyllum and 
Funis (see Woelkerling, 1972). The specimens have been 
referred to Elachistea lubrica because of the apparent ab- 
sence of moniliform paraphyses (see Taylor, 1957, p. 139), 






1975] Sargasso Sea — Woelkerling 13 



determine 



reliable character of specific distinction. Lund (1959) and 
Rosenvinge (1893) have regarded E. lubrica as a variety 



/ 



Family Chordariaceae 
Genus Chordaria C. Agardh, 1817 



Chordaria flagelliformis (Mueller) C. Agardh. Kylin 1947: 
59, Figs. 51A, D. Lund 1959:121, Figs. 26, 27. Taylor 
1957 : 148, pi. 12, Fig. 6 ; pi. 14, Fig. 4. 
type locality: Denmark. 

TYPE : not located. 

distribution: cooler waters of North America and Eu- 
rope. 

specimens examinrd: Sargasso Sea: 39°05'N-69°48'W, lO.v.1970, 
Woelkerling (WJW 2576) ; 39°11'N-69°24'W, lO.v.1970, Woelkerling 
(WJW 2554). 

The plants were epiphytes on invading specimens of 
Fucus (see Woelkerling, 1972) and are not considered per- 
manent components of the Sargasso Sea flora. 



ORDER PUNCTARIALES 

Family Punctariaceae 
Genus Colpomenia Derbes and Solier, 1856 



mem 



110, pi. 7, Fig. 1, pi. 19, Figs. 3-4; 1960 :260, pi. 36, Fig. 1. 

Womersley 1967 :244. 

type locality : Cadiz, Spain. 

type : probably lost. 

distribution : widespread. 

specimen examined: Sargasso Sea: 32°09'N-64°58'W, 16.V.1970, 
Woelkerling (WJW 2655). 



i almost cert, 
om Bermuda 



permanent com 



14 Rhodora [Vol. 77 






Genus Petalonia Derbes and Solier, 1850 
Petalonia fascia (Mueller) Kuntze. Lund 1947:31, Fig 10 
Taylor 1957:167, pi. 14, Fig. 5; pl.15, Fig. 3. 

Ilea fascia (Mueller) Fries. Kylin 1947:77, Fig. 61A. 
type locality: Denmark. 
type : not located. 
distribution : widespread. 

specimen examined: Sargasso Sea: 39°11'N-69°24'W 10 v 1970 

Woetteerling (WJW 2549). 

This taxon is an invading element attached to Fucus 
vcsiculosus (see Woelkerling, 1972). 

Genus Punctaria Greville, 1830 
Punctaria latifolia Greville. Collins, Holden, and Setchell 

1895:82; 1901:873; 1907:1388. Taylor 1957:166, pi 15 
Fig. 5. 

type locality : Great Britain. 
type: not located. 
distribution i widespread. 

specimen examined: Sargasso Sea: 39°0.VN-69°48'W 10 v 1970 
Woelkerling (WJW 2573). 

A single, rather small and battered plant was found at- 
tached to an invading Fucus element (see Woelkerling 
1972). 

F'unctaria plantaginea (Roth) Greville. Rosenvinge et Lund 
1947:11, Fig. 2; 1959:133, Fig. 28. Taylor 1957:166, pi. 
15, Fig. 4; pi. 16, Fig. 4. 

type locality : Kattegat Channel between Denmark and 
Sweden. 

TYPE : probably destroyed. 
distribution : widespread. 

specimens examined: Sargasso Sea: 38 o 53'N-09°39'W, 10 v 1970 
Woelkerling (WJW 2560) ; 39 o 05'N-(59 o 48'W, lO.v.1970, Woe'lk'erHng 
(WJW 2575). 

Two small plants occurred as epiphytes on Fucus; they 
are not considered permanent components of the Sargasso 
Sea flora (see Woelkerline-. 1972K 



1975] Sargasso Sea — Woelkerling 15 



Genus Scytosiphon C. Agardh, 1811 

:ytosiphon lomentaria (Lyngbye) C. Agardh. Rosenvinge 
et Lund 1947:27, Fig. 9; 1959:103, Fig. 20. Taylor 1957: 
168, pi. 15, Fig. 2 ; pi. 16, Fig. 3. 

type locality: Denmark. 

type: C. 

distribution: nearly cosmopolitan. 

specimens examined: Sargasso Sea: 38°53'N-69°39'W, lO.v.1970, 



Wo 



Wo 



(WJW 2574) ; 39°11'N-69°24'W, lO.v.1970, Woelkerling (WJW 2548). 

These collections occurred as epiphytes on Fucus and 
probably do not represent permanent components of the 
Sargasso Sea flora. The species does, however, occur in 

and along the southeastern U. S. Coast (Taylor, 
d may eventually be found to occur on Sargassum 



Bermuda 



fluitans 



Family Striariaceae 

Genus Isthmoplea Kjellman, 1877 

Isthmoplea sphaerophora (Harvey in Hooker) Kjellman. 
DeToni 1895:569. Kylin 1947:67, Figs. 56D-E. Taylor 
1957:156, pi. 9, Figs. 4-5. Woelkerling 1972:298. 

Ectocarpus sphaerophorus Carmichael. Harvey 1846: 

pi. CXXVI. 
TYPE locality : Appin, Scotland. 

type: tcd. 

distribution: cooler waters of Eastern North America 
and Europe. 

specimen examined: Sargasso Sea: 38°53'N-69°39'W, lO.v.1970, 
Woelkerling (WJW 2566). 



The 



3 



mb 



attached to Polysiphonia lanosa, in turn an epiphyte on a 
plant of Ascophyllum, which had drifted out into the 
Northwestern fringes of the Sargasso Sea (see Woelker- 
ling, 1972). 



16 Rhodora [Vol. 77 



ORDER FUCALES 

Family Fucaceae 

Genus Ascophyllum Stackhouse 1809 

Ascophy Hum nodosum (L.) Le Jolis. Kylin 1947:84. Taylor 
1957:195, pi. 27, Figs. 1-2. 

type locality : Atlantic Ocean. 

TYPE: LINN. 

distribution : widespread in colder waters of the north- 
ern hemisphere. 

specimens examined: Sargasso Sea: 38°53'N-f>9°39'W, lO.v.1970, 
Woelkerling (WJW 2558) ; 39°05'N-69°48'W, lO.v.1970, Woelkerling 
(WJW 2568) ; 39°11'N-69°24'W, lO.v.1970, Woelkerling (WJW 2546). 

The plants were found adrift along the northwest fringes 
of the Sargasso Sea; there is some question as to whether 
they should be regarded as permanent components of the 
Sargasso Sea flora (see Woelkerling, 1972). 

Genus Fucus Linneaus, 1753 

Fucus vesiculosus L. Harvey 1852:71. Kylin 1947:83, Tab. 
17, Figs. 53-54. Taylor 1957:192, pi. 25, Figs. 1-3. 

type locality: Atlantic Ocean. 

type: linn. 

distribution: widespread in colder waters of northern 
hemisphere. 

specimens examined: Sargasso Sea: 38°53'N-69°39'W, lO.v.1970, 
Woelkerling (WJW 2563): 39°05'N-69°4S 



W. lO.v.1970. Wo 



(WJW 2570); 39°11'N-69°24'W, lO.v.1970, Woelkerling (WJW 



were 



of the Sargasso Sea with Ascophyllum and Sargassum 
natans and bore a number of epiphytes (see Woelkerling, 



1972). 



Family Sargassaceae 
Genus Sargassum C. Agardh, 1820 



The tropical American Atlantic species of Sargassum are 
poorly known and specific limits and distinctions remain 



1975] Sargasso Sea — Woelkerling 17 



very unclear (see Taylor, 1960, p. 268). Moreover, keys to 
species occurring in this region (e.g. Howe, 1920; Taylor, 
1928, 1960) are based largely on vegetative features rather 



more 



fundamental significance (Setchell, 1931 ; Womersley, 1954). 
The situation is further complicated by the fact that the 
two species of Sargassum most commonly found in the Sar- 
gasso Sea apparently have never been found in a fertile 
state (see, however, Parr, 1939). 

During the course of this study, seven apparently distinct 
species of Sargassum have been found adrift in the western 
Sargasso Sea. Only three of these, however, have been 
identified with any certainty. The remaining four have not 
been definitely identified as yet and, following the approach 
of Winge (1923), are referred to here as Sargassum A, B, 

- — M m 



name confusion until such time 



mono 



this region. 



Differences between the seven taxa are summarized as 



follows: 



Conspectus of Sargassum Taxa in 
the Western Sargasso Sea 



1. Plants sterile, pelagic, without evidence of a basal hold- 



fast 



2. 



2. Stems smooth ; vesicles often apiculate ; leaves linear, 
up to 4 mm wide £• natans. 

2. Stems muriculate ; vesicles at most muticous ; leaves 
lanceolate, up to 8 mm wide S. fluitans. 

1. Plants commonly fertile, normally attached, usually 

showing evidence of a holdfast 3. 

3. Fruiting branches often carpophyllaceous (i.e. of 

mixed receptacles, leaves, and vesicles) 4. 

4. Receptacles pedicellate ; cryptostomata tending to 

be in a single row adjacent to the costa 

"Sargassum D." 



18 Rhodora [Vol. 77 



4. Receptacles not pedicellate; cryptostomata scat- 
tered "Sargassum B." 

3. Fruiting- branches not carpophyllaceous (i.e. com- 
posed solely of receptacles) 5. 

5. Receptacles pedicellate 6. 

6. Receptacles commonly spiny; costa not spiny 

or dentate 

. Sargassum hystrix var. buxi folium. 
6. Receptacles not spiny; costa prominently den- 
tate to spiny "Sargassum A." 

5. Receptacles sessile "Sargassum C." 

Sargassum fluitans (Boergesen) Boergesen 1914a :6. Taylor 
1928:127, pi. 18, Fig. 9; pi. 19, Fig. 5; 1960:281, pi. 39, 
Fig. 2, pi. 40, Fig. 7. 

Sargassum hystrix J. Ag. var. fluitans Boergesen 1914a: 
11, Fig. 8. Winge 1923:23, Fig. 6 (as "Sargassum 
III"). 

TYPE locality : Sargasso Sea. 
TYPE: c. 

distribution: known only from the Sargasso Sea. 

specimens examined: Sargasso Sea: 21°58'N-68°20'W, 24.iv.1970, 
Moore (WJW 2639) ; 26°57'N-72°58'W, 26.iv.1970, Moore (WJW ' 
2649); 28°N-70°W, 4.vii.l970, Volkmanv (WJW 2738); 31°N-69° 
29'W, 3.iii.l970, Volkmann (WJW 2394); 31°N-70°W,' 5.vii.l970, 
Volkmann (WJW 2731); 32°09'N-64°58'W, 16.V.1970, WoelkerKng 
(WJW 2654) ; 33°58'N-69°56'W, 15.V.1970, Woelkerling (WJW 2602); 
34°N-70°W, lO.i.1970, Volkmavtn (WJW 2224), 7.iii.l970, Volkmann 

(WJW 2418), 6.vii.l970, Volkmann (WJW 2742), 14.viii.1970, Moore 
(WJW 2933); 36°N-70°36'W, 9.xii.l970, Moore (WJW 2947); 37°N- 
70°W, 12.V.1970, WoelkerKng (WJW 2616); 37°30'N-70°W, 8.vii.l970, 
Volkmann (WJW 2712); 38°34'N-09°11'W, 19.V.1970, Woelkerling 
(WJW 2606). 

Sargassum, fluitans is the less frequently encountered of 
the two species of Sargassum endemic to the Sargasso Sea 
but apparently enjoys almost as wide a distribution. The 
specimens examined during this study were commonly cov- 
ered with bryozoans and to a lesser extent with epibiotic 
algae. 



1975] Sargasso Sea — Woelkerling 19 



Sargassum hystrix J. Agardh var. buxifolium (Chauvin) 

J. Agardh 1889:91, tab VII, Fig. 1. Boergesen 1914:221. 

Chapman 1963:45. Earle 1969:225, Fig. 118. Grunow 

1915:399. Howe 1920:594. Taylor 1928:128, pi. 18, 

Fig. 1, pi. 19, Fig. 9; 1960:279, pi. 38, Fig. 2, pi. 40, 
Fig. 6. 

type locality: Caribbean Area (see DeToni 1895:53). 
type: ld(?). 

distribution : Florida to Brazil, Caribbean Islands, Sar- 
gasso Sea. 

specimens examined: Sargasso Sea: 36°N-70°36'W, 9.xii.l!)70, 



Moore (WJW 2912); 39°30'N-71°W, 6.X.1970, Volkmann (WJW 
2873). 

The specimens examined agree well with the descriptions 
and illustrations of Earle (1969) and Taylor (1960), and 
probably are the same as "Scvrgassum VI" of Winge (1923, 
p. 25, Fig. 10). The material was collected in the immediate 
vicinity of the Gulf Stream, the same region reported by 
Winge (1923), and apparently is known mainly from drift 
specimens (Earle, 1969; Taylor, 1928, 1960). 

Both specimens bore non-carpophyllaceous receptacles 
which were simple or once funcate, terete or slightly com- 
pressed, verrucose or occasionally with odd spines, pedi- 
cellate, and more or less racemose. 

Sargassum natans L. Boergesen 1914a :7, Figs. 3-7. Taylor 
1928:128, pi. 18, Figs. 2-4, pi. 19, Fig. 13; 1960: pi. 37, 
Fig. 2, pi. 40, Figs. 3, 8. Winge 1923:24, Figs. 3-5 (as 
"Sargassum I & III"). 
TYPE locality : Sargasso Sea. 

type: linn. 

distribution : known only from the Sargasso Sea. 



specimens examined: Sargasso Sea: 20°f)0'N-67°15'W, 24.iv.1970, 
Moore (WJW 2645); 21 o 58'N-68°20'W, 24.iv.1970, Moore (WJW 
2640); 26°50'N-71°48'W, 5.iii.l970, Volkmann (WJW 2422); 26°57'N- 
72°58'W, 26.iv.1970, Moore (WJW 2653); 28°N-70°W, 4.iii.l970, 
Volkmann (WJW 2437). 4.vii.l970, Volkmann (WJW 2739), 31°N- 
69°29'W, 3.iii.l970, Volkmann (WJW 2393); 31°N-70°W, 5.vii.l970, 
Volkmann (WJW 2732); 32°09'N-64°58'W, 16.V.1970, Woelkcrlinf/ 
(WJW" 2662); 34°N-70°W, lO.i.1970, Volkma'nn (WJW 2223), 7.iii. 



20 Rhodora [Vol. 77 



1970, Volkmann (WJW 2417), 13.V.1970, Volkmann (WJW 2607), 
30.vi.1970, Volkmann (WJW 2723), 6.vii.l970, Volkmann (WJW 
2743), 14.viii.1970, Volkmann (WJW 2931); 35°54'N-70°30'W, 13.viii. 



1970, Moore (WJW 



JW 



2950); SO^'N^O^'W, 15.viii.1970, Moore (WJW 2905); 37°N- 
70°W, 12.V.1970, Woelkerling (WJW 2617) ; 37°30'N-70°W, 8.vii.l970, 
Woelkerling (WJW 2711); 38°22'N-70°58'W, 12.X.1970, Volkmann 
(WJW 2897) ; 38°34'N-69°11'W, 19.V.1970, Woelkerling (WJW 2605) ; 
39°07'N-70°35'W, 16.viii.1970, Moore (WJW 2909); 39°11'N-69°24'W, 
lO.v.1970, Woelkerling (WJW 2644). 

Sargassum natans plants comprise the vast bulk of the 
Sargasso Sea macroscopic vegetation, estimated by Parr 
(1939) to be up to 40 million metric tons. It also harbored 
the greatest variety and quantity of epibiotic algae. The 
considerable quantities of S. natans which wash up on the 
shores of Bermuda, in contrast, are apparently devoid of 
epiphytes (Woelkerling, personal observations). 

"Sargassum A" 

The single specimen [39°30'N-70°W, 6.x. 1970, Volkmann 
(WJW 2867)] referred to this "taxon" bears a small, dis- 
coid holdfast and a sparsely branched, nearly terete main 
axis with a few, scattered, long laterals whose stems are 
muriculate. The leaves are lanceolate, up to 3 mm broad 
and 30 mm long, finely serrate, with a prominent dentate to 
spiny costa. Cryptostomata are lacking. Vesicles up to 5 
mm in diameter are mostly single and pedicellate and scat- 
tered among the leaves. 



simpl 



verrucose 



mm 



While these features are most closely associated with 
Sargassum filipendula var. montagnei as described in Taylor 
(1960), definite specific affiliation of the specimen in ques- 
tion remains uncertain. 

"Sargassum B" 

The single specimen [34°N-70°W, lO.i.1970, Volkmann 
(WJW 2222)] lacks a holdfast and has a smooth stem with 
a number of short lateral branches. The leaves are mostly 



1975] Sargasso Sea — Woelkerling 21 



lanceolate ovate, up to 5 mm broad and 30 mm long, serrate, 
have costae without ridges or wings and bear scattered 
cryptostomata. Vesicles up to 6 mm broad are short pedi- 
cellate and are scattered among the leaves and receptacles. 

The receptacles are generally carpophyllaceous and usu- 
ally 2-4 times furcate, are terete, not dentate or spiny, are 
not pedicellate and are borne in dense cymose clusters. 

Although this taxon shows many of the features of Sar- 
gassum vulgare C. Agardh as described in Taylor (1960), 
final specific identification remains uncertain. 

"Sargassum C" 

The single specimen [33°58.5'N-69°56.5'W, 15.V.1970, 
Woelkerling (WJW 2589)] lacks a holdfast and has mu- 
riculate stems bearing lateral branches of variable length. 
The leaves are linear lanceolate, up to 3 mm broad and 30 
mm long, entire or finely serrate, and bear inconspicuous 
costae and scattered cryptostomata. Shortly pedicellate 
vesicles up to 5 mm in diameter are scattered along the 

branches. 

The receptacles generally are not carpophyllaceous, are 
one to several times furcate, terete, verrucose, generally 
not pedicellate, and are racemose. 

This specimen could not be linked to any of the species 
described by Taylor (1960). In some respects it is similar 
to what Taylor (1960) calls Sargassum filipendula var. 
montagnei, but differs in the nature of the costa and in 
having non-pedicellate receptacles. 

"Sargassum D" 

The one specimen [34°N-70°W, lO.i.1970, Volkmann 
(WJW 2225)] lacks a holdfast and has a stem that is mu- 
riculate in the younger portions and smooth in the older 
portions. Lateral branches vary in length and bear lanceo- 
late leaves up to 3 mm broad and 30 mm long which are 
mostly entire, costate, and bear cryptostomata which tend 
to lie in a single row on each side of the costa. Shortly 
pedicellate vesicles up to 5 mm in diameter are scattered 

along the lateral branches. 



22 



Rhodora [Vol. 77 



The receptacles are carpophyllaceous, simple or once 
furcate, terete, not dentate or spiny, pedicellate, and gen- 
erally cymose. 

Of the species described by Taylor (1960), this specimen 
most closely approximates Sargasswm acinarium (L.) C. 
Agardh, but apparent differences in receptacle morphology 
leaves some doubt as to the specimen's true affinities. 

A second specimen [37°N-70°W, 12.V.1970, Woelkerling 
(WJW 2673)] shares many features with "Sargassum D" 
including the linear distribution of cryptostomata, but the 
racemose nature of the receptacles leaves some doubt as to 
its exact relationships until further material becomes avail- 
able for study. 

CLASS RHODOPHYCEAE 

Subclass Bangiophycidae 

order bangiales 

Family Bangiaceae 

Genus Asterocytis Gobi, 1879 

Asterocytis ramosa (Twaites in Harvey) Gobi. Boergesen 
1915:3, Fig. 1. Chapman 1963:49. Kylin 1944:6, Fig. 
ld-f. Rosenvinge 1909:77, Fig. 17. Taylor 1928:132, 
pi. 20, Figs. 1-2; 1960:287. Wille 1900:7, Tab. 1, Figs. 
8-14. 

Hormospora ramosa Twaites in Harvey 1846-51 :P1. 

CCXII. 

TYPE locality: Wareham, Dorsetshire, Great Britain. 

TYPE: TCD. 

distribution : widespread. 

SPECIMENS EXAMINED: Sargasso Sea: 37°N-70°W, 12.V.1970, Woel- 
kerling (WJW 3957); 39°07'N-70°35'W, 16.viii.1970, Moore (WJW 
2920). 

In both cases the plants were epiphytic on Ckidophora 
which, in turn, was growing on Sargassum nutans. 
Hamel (1924), Boergesen (1927), and Pham 



(1969), among others, have referred this species to the 
synonomy of Asterocystis ornata (C. Agardh) Hamel. 



1975] Sargasso Sea — Woelkerling 23 



Kylin (1944) and Taylor (1957, 1960), however, prefer to 



ramosa is marine 



and A. ornata freshwater. The validity of such a separation 
requires further investigation, and until new evidence 
comes to light, the two taxa will be kept distinct for pur- 
poses of the present study. 

Genus Erythrocladia Rosenvinge, 1909 

Erythrocladia subintegra Rosenvinge 1909:73, Figs. 13-14. 
Boergesen 1915:7, Figs. 3-4. Collins and Hervey 1917:95. 






Pham-Hoang-Ho- 1969 :80, Figs. 2-12. Taylor 1960:290. 
TYPE locality: Hirshals, Skagerak, Denmark. 
TYPE: c. 

distribution: Europe, western tropical Atlantic, South- 
east Asia. 

SPECIMENS examined: Sargasso Sea: 31°N-69°29'W, 3.iii.l970, 
Volkmann (WJW 2387, 2391, 2392) ; 34°N-70°W, 10.L1970, Volfc- 
malnn (WJW 2212), 7.iii.l970, Volkmann (WJW 2405, 2406); 36°N- 
70°36'W, 9.xii.l970, Moore (WJW 2939, 2940). 

All specimens occurred as epiphytes on hydroids which, 



fl 



gassum 



Erythrocladia recondita Howe et Hoyt 1916:112, pi. 12, 

Figs. 1-5, pi. 13, Fig. 1. Hoyt 1920:467, pi. CXVI, Fig. 1, 

pi. CXVII, Figs. 1-5. 

TYPE locality : Beaufort, North Carolina. 
type: NY. 

distribution : type locality, Sargasso Sea. 

specimen examined: Sargasso Sea: 39°30'N-71°W, 6.x. 1970, Volk- 
mann (WJW 2866). 

The specimens occurred epizoically on hydroids which in 
turn grew on an unidentified Sargassum. 

Genus Erythrotrichia Areschoug, 1850 

Erythrotrichia carnea (Dillwyn) J. Agardh. Boergesen 
1915:7. Collins and Hervey 1917:94. Hoyt 1920:466, 
Fig. 24. Rosenvinge 1909:67, Fig. 8. Taylor 1957:202, 
pi. 28, Figs. 13-15 ; 1960 :292. Woelkerling 1972 :298. 



24 



Rhodora [Vol. 77 



type locality : Great Britain. 
type: nmw. 
distribution : widespread. 

specimens examined: Sargasso Sea: 28°N-70°W, 4.iii.l970, Volk- 
mann (WJW 2433); 31°N-69°29'W, 3.iii.l970, Volkmami (WJW 
2360; 2390); 34°N-70°\V, lO.i.1970, Volkmaim (WJW 2216), 7.iii.l970, 
Volkmann (WJW 2404); 39°11'N-69°24'W, lO.v.1970, Woelkerling 
(WJW 2551). 

With the exception of the last cited specimen, all plants 
occurred epizoically on hydroids which in turn were at- 
tached to Sargassum fliu'tans, S. natans, or Sargassum sp. 
In the other collection, the plants occurred epiphytically on 
Fucus (Woelkerling, 1972). 

Subclass Florideophycidae 

order nemaliales 

Family Acrochaetiaceae 

Four species of Andouinella (A. daviesii, A. hallandica, 
A. microscopica, A. saviana [= A. thurctii]) and two spe- 
cies of Colaconema (C. infestans, C. secundata) occur in 
the western Sargasso Sea and are the subject of a recent 
detailed morphotaxonomic study (Woelkerling, 1973). 

ORDER CRYPTONEMIALES 

Family Corallinaceae 
Subfamily Corallineae 

Genus Jania Lamouroux, 1812 

Jania adherens Lamouroux. Boergesen 1917:195, Figs. 184- 
187. Chapman 1963:86, Fig. 85. Taylor 1928:205. Howe 
1920:589. 1960:413, pi. 49, Figs. 1-2. 
type locality : Mediterranean Sea. 
type : not located. 

distribution: widespread in tropical and warm temper- 
ate waters. 

specimen examined: Sargasso Sea: 32°09'N-65°58'W, 16.V.1970, 
Woelkerling (WJW 2656). 

The single collection occurred as an epiphyte on a plant 
of Colpomenm (q.v.) which probably drifted out from the 



1975] Sargasso Sea — Woelkerling 25 



Bermuda Islands; consequently this species of Jania prob- 
ably does not represent a permanent component of the Sar- 
gasso Sea flora. 

Jania capillacea Harvey 1853:85. Boergesen 1917:198, Fig. 

188. Chapman 1963:86, Fig. 86. Collins, Holden, and 

Setchell 1895:150. Howe 1920:589. Taylor 1928:206, 

pi. 29, Figs. 2, 10. 1960:413, pi. 49, Figs. 1-2. 

type locality : Bahia Honda, Florida. 

type: tcd. 

distribution: tropical western Atlantic. 

specimens examined: Sargasso Sea: 31°N-69°29'\V, 3. Hi. 1970, 
Volkmann (WJW 2217); 36°N-70°36'W, 9.xii.l970, Moore (WJW 
2945); 39°07'N-70°35'W, 16.viii.1970, Moore (WJW 2926). 

Jania capillacea has been found growing on Sargassum 
fluitans, S. natans, and on several unidentified Sargassum 



taxa. Prat (1935) 
Sargasso Sea. 



from 



Subfamily Melobesieae 

Genus Fosliella Howe, 1920 

Fosliella farinosa (Lamouroux) Howe 1920:587. Chapman 
1963:91, Fig. 92. Dawson 1960:30, pi. 21, Fig. 1, pi. 22, 
Fig. 1. Taylor 1960:388. Womersley and Bailey 1970: 

309. 

Melobesia farinosa Lamouroux. Lemoine in Boergesen 

1917:170, Fig. 165. Hoyt 1920:523, Fig. 47. Taylor 

1928:211. 
type locality : Adriatic Sea. 

type: cn. 

distribution: nearly cosmopolitan. 

specimens examined: Sargasso Sea: 31°N-69°29'W, 3.iii.l970, 
Volkmann (WJW 2355); 34°N-70°W, 13.V.1970, Woelkerling (WJW 
2608). 

The plants occurred epiphytically on Sargassum fluitans 
and S. natans. 

Fosiella lejolisii (Rosanoff) Howe 1920:588. Masaki 1968: 
23, pis. XII, XLIX, L. Taylor 1957:253, pi. 36, Figs. 6-8. 



26 Rhodora [Vol. 77 



Melobesm lejolisii Rosanoff 1866:62, pi. 1, Figs. 1-13, pi. 
7, Figs. 9-11. Rosenvinge 1917:238, Figs. 156-159. 

type locality : Cherbourg, France. 

type : not located. 

distribution: Europe, Atlantic North America, Japan, 



Sargasso Sea. 



KX 



Sargasso Sea: 28°N-70°W, 4.iii.l970, Volk- 



mann (WJW 2436): 31°N-69°29'W 



2378); 32°09'N-64°58'W. 16.V.1970, Woelkerling (WJW 2659, 2672); 
33°58.5'N-69°56.5'W, 15.V.1970, Woelkerling (WJW 2590, 2600); 



WJW 



9.xii.l970, Moore (WJW 2946 



Woel 



kerling (WJW 2634); 39°07'N-70°35'W, 16.viii.1970, Moore (WJW 
2928). 

Fosliella lejolesii occurs as an epiphyte on Sargassum 

fluitans, S. natans, Sargassum sp. as well as on other algae, 

and in terms of numbers of individuals, is probably the 

most common red alga in the Sargasso Sea. Only Ceramium 

gracillimum (q.v.) approaches the same quantitative fre- 
quency. 

According to Taylor (1960, p. 387) Fosiella lejolisii is 



from L. affi 



um 



thallus cells 6-7 /xm broad rather than 9-18 

10-12 ^m broad. However, cells up to 13 jxm broad were 

found in Sargasso Sea collections, and this suggests that 

the relationships of the three taxa require critical reinvesti- 
gation. 



ORDER CERAMIALES 

Family Ceramiaceae 

Genus Awtithamnion Naegeli, 1847 

Antithamnion a nf ilia i u in Boergesen 1917 :226, Figs. 213-216. 
Taylor 1960:499. Womersley and Bailey 1970:322. 

type locality : St. Thomas, Virgin Islands. 
type: c. 

distribution: Virgin Islands, Sargasso Sea, tropical 
Pacific Ocean. 



1975] Sargasso Sea — Woelkerling 27 



specimens examined: Sargasso Sea: 28°N-70°W, 4.iii.l970, Volk- 
mann (WJW 2426); 31°N-69°29'W, 3.iii.l970, Volkmann (WJW 



2376). 



phytically on Sargassum 



natum. One (WJW 2376) bore tetrasporangia ; the other 
was sterile. The genus Antithamnion in this area needs 
critical reinvestigation in light of recent studies of Wollas- 
ton (1968, 1971) on southern Australian and Pacific Coast 
species. 

Genus Ceramium Roth, 1797 

Ceramium gracillimum (Kuetzing) Griffiths and Harvey. 

Dawson 1962:57, pi. 20, Figs. 2-3. Feldmann-Mazoyer 

in Boergesen 1952:42, Fig. 21. Nakamura 1965:136, 

pi. 1, 5-6, Fig. 6. 

Ceramium gracillimum var. byssoideum (Harvey) Ma- 
zoyer 1938:323. Chapman 1963:178, Fig. 186. Feld- 
mann-Mazoyer 1940:293, Fig. 109. 

Ceramium byssoideum Harvey 1853:218. Taylor 1928: 
190, pi. 27, Figs. 20, 21; 1960:528, pi. 67, Figs. 1-3. 

Ceramium transversale Collins and Hervey 1917:145, 

pi. 5, Figs. 29-31. Boergesen 1918:243. 
TYPE LOCALITY I Trieste, Italy. 



type: c. 

distribution : wides 
ate seas ; England. 



temper- 



KX 



Moore (WJW 2642); 26°50'N-71°48'W, 5.iii.l970, Volkmann (WJW 
2419); 26°57'N-72 58'W, 26.iv.1970, Woelkerling (WJW 2652); 28°N- 
70°W, 4.iii.l970, Volkmami (WJW 2428); 31°N-70°W, 5.vii.l970, 
Volkmann (WJW 2733); 32°09'N-64°58'W, 16.V.1970, Woelkerling 
(WJW 2661, 2669) ; 33°58.5'N-69°56.5'W, 16.V.1970, Woelkerling 
(WJW 2591)'; 34°N-70°W, 13.V.1970, Woelkerling (WJW 2609), 6.vii. 
1970, Volkmann (WJW 2746) ; 35°54'N-70°30'W, 13.viii.1970, Moore 
(WJW 2903); 37°N-70°W, 12.V.1970, Woelkerling (WJW 2619); 
37°30'N-70°W, 8.vii.l970. Volkmann (WJW 2705); 38°22'N-70°58'W, 
12.X.1970, Volkmann (WJW 2890). 

Ceramium gracillimum is a common epiphyte on Sargas- 
sum fluitans, S. natans, and Sargassum sp. as well as on 



28 Rhodora [Vol. 77 



other algae attached to Sargassum and often occurs in 
considerable numbers. One collection (WJW 2619) bore 
tetrasporangial individuals ; the remainder were sterile. 

Taylor (1960, p. 528) maintains Ceramium byssoideum 
and C. gracillimum as distinct taxa; however, following 
Feldmann-Mazoyer (1940), Nakamura (1965) and others 
the former is referred to the conspecificity of the latter. 
Likewise, C. transvcrsale is regarded as conspecific in ac- 
cordance with Feldmann-Mazoyer (1940). 

Ceramium fastigiatum Harvey in Hooker [non C. fastigi- 
atum Roth = Polysiphonia fastigiata (Roth) Greville]. 

Boergesen 1918:241, Fig. 231. Chapman 1963:177, Fig. 

184a-c. Nakamura 1965:129, pi. 1(3), Fig. 4. Taylor 

1928:191; 1957:309, pi. 47, Figs. 3-5, 7, pi. 48, Figs. 2-4, 

pi. 49, Figs. 3-4, pi. 50, Fig. 4, pi. 51, Figs. 6-7; 1960:526, 

pi. 67, Figs. 4-6. 

TYPE locality: Great Britain. 

type: tcd. 

distribution : widespread. 

specimens examined: Sargasso Sea: 28°N-70°W, 4. Hi. 1970, Volk- 
mann (WJW 2427) ; 31°N-69°29'W, 3.iii.l970, Volkmann (WJW 2348, 
2370, 2373) ; 34°N-70°W, 7.iii.l970, Volkmann (WJW 2395); 39°07'N- 
70°35'W, 16.viii.1970, Moore (WJW 2923). 

This taxon occurs as an epiphyte on Sargassum fluitans, 
S. natans, and Sargassum sp. All specimens examined were 
sterile. The author citations of Taylor (1960) and Boerge- 
sen (1918) (i.e. C. f astig iatium (Roth) Harvey) are in- 
correct; Roth's taxon was described from Germany and 
has been referred by Greville (1824) to Polysiphonia (see 
DeToni, 1903, p. 945-6) whereas Harvey's taxon was de- 
scribed from Great Britain (Harvey in Hooker, 1833). 

Genus Crouania J. Agardh, 1842 

Crouania attenuata (C. Agardh) J. Agardh, Boergesen 
1917:230, Figs. 219-221. Chapman 1963:167, Fig. 173. 
Collins and Hervey 1917:142. Harvey 1853:226, Tab. 
XXXI, D. Taylor 1928:193, pi. 27, Figs. 7-9, pi. 32, 
Fig. 9. 



1975] Sargasso Sea — Woelkerling 29 



type locality : Mediterranean Sea. 

type: ld. 

distribution: Mediterranean, England, tropical western 

Atlantic Ocean, Japan. 

specimens examined: Sargasso Sea: 37°N-70°W, 12.V.1970, Woel- 
kerling (WJW 2626); 39°30'N-71°W, 6.X.1970, Volkmann (WJW 
2882). 

Male (WJW 2626) and tetrasporangial (WJW 2882) 



piphytes on Sargassum 



gassum 



Genus Griffithsia C. Agardh, 1817 

Griffithsia radicans Kuetzing 1862:11, tab. 33, Fig. A-C. 
Taylor 1960:515. 
type locality: Brazil. 
type: l. 

DISTRIBUTION : Brazil, Sargasso Sea. 

specimens examined: Sargasso Sea: 37°N-70°W, 12.V.1970, Woel- 
kerling (WJW 2629); 39 o 30'N-71°W, 6.X.1970, Volkmann (WJW 



2881). 



epiphytes on Sargassum natans 



qassiim sp. and appeared to have some 



gial initials. They are referred to this species because of 
their agreement with the description given by Taylor 
(1960). 

Genus Spermothamnion Areschoug, 1847 



mothamnion 



Maze et Schramm 



Vickers. Boergesen 1909:17, Fig. 10; 1917:200, Figs. 189- 
190; 1920:461, Fig. 422. Collins and Hervey 1917:132. 
Howe 1920 :578. Taylor 1960 :520. 
type locality: Guadeloupe. 

type: PC. 

distribution: North Carolina, Caribbean Islands, Sar- 
gasso Sea. 

specimens examined: Sargasso Sea: 39°30'N-71°W, 6.X.1970, 
Volkmann (WJW 2863). 



em 



de and tetrasporangial plants were found as epi- 
phytes on "Sargassum A" and may not, therefore, be a 



30 Rhodora [Vol. 77 



permanent component 



Tropical 



and subtropical western Atlantic members of the Sper- 
mothamneae are in need of thorough reinvestigation in 
light of the recent study of Gordon (1972). 

Genus Wrung elia C. Agardh, 1828 

Wrangelia argus (Montagne) Montage. Boergesen 1916: 
116, Figs. 125-6. Gordon 1972:40. Taylor 1928:144, pi. 
20, Fig. 13, pi. 22, Fig. 6, pi. 32, Fig. 4; 1960:502, pi. 66, 

Figs. 7-8. 

TYPE locality: unknown. 
type : probably c. 

distribution: see Gordon 1972, p. 40. 

specimens examined: Sargasso Sea: 31°N-69°29'W, 3.iii.l970, 
Volkmann (WJW 2371); 37°N-70°W, 12.V.1970, Woelkerling (WJW 
2628); 39°30'N-71°W, 6.X.1970, Volkmann (WJW 2880). 

Male and tetrasporangial plants occurred as epiphytes 
on Sargassum natans and on Sargassum sp. 

Family Dasyaceae 

Genus Dasya C. Agardh, 1824 

Dasya rigidula (Kuetzing) Ardissone. Howe 1920:576. 

Taylor 1960 :558, pi. 72, Fig. 4. 

type locality : Spalato, Adriatic Sea. 
type: l. 

distribution : Bermuda, Caribbean Islands, Mexico, Sar- 
gasso Sea, Venezuela, Adriatic and Mediterranean Seas. 

specimens examined: Sargasso Sea: 34°N-70°W, 7.iii.l970, Volk- 
mann (WJW 2398); 36°N-70 o 36'W, 9.xii.l970, Moore (WJW 2941); 
39°30'N-71°W, fi.x.1970, Volkmann (WJW 2861, 2884). 

Male, female, and tetrasporangial plants occurred epi- 
phytically on Sargassum fluitans and on Sargassum sp. 

Genus Heterosiphonia Montagne, 1842 

Heterosiphonia wurdemanni (Bailey in Harvey) Falkenberg 
1901:638, pi. 16, Fig. 11. Boergesen 1919:324, Figs. 326- 
328. Collins and Hervey 1917:131. Howe 1920:575. 
Taylor 1928:178, pi. 25, Fig. 3; 1960:565, pi. 72, Fig. 9. 



1975] Sargasso Sea — Woelkerling 31 



Dasija wurdemanni Bailey in Harvey 1853:64, Tab. XV, 

C. 

type locality : Key West, Florida. 

type: tcd. 

distribution: western tropical Atlantic; Mediterranean. 

specimens examined: Sargasso Sea: 34°N-70°W, lO.i.1970, Volk- 
mann (WJW 2213). 

A single tetrasporangial plant occurred epiphytically on 
"Sargassum B"; its status in the Sargasso Sea flora must 
remain in doubt until further collections come to hand. 

Family Delesseriaceae 

Genus Hypoglossum Kuetzing, 1843 

Hypoglossum tenuifolium (Harvey) J. Agardh. Howe 1920: 
564. Taylor 1960 :545, pi. 68, Fig. 2. 

Delesseria tenuifolia Harvey 1853:97, Tab. XXII, Fig. B. 

Boergesen 1919:344, Figs. 340-343. 
type LOCALITY : Key West, Florida. 

TYPE: TCD. 

distribution: western tropical Atlantic Ocean. 

specimens examined: Sargasso Sea: 34°N-70°W, 10. i. 1970, Volk- 



(WJW 



ant was found epiphytically on "Sargaasi 
further collections come to hand, its statu* 
comnonent of the Sargasso Sea flora must 



main in doubt. 



Family Rhodomelaceae 



Genus Chondria C. Agardh, 1817 

Two collections — one tetrasporangial [37°N-70°W, 
12.V.1970, Woelkerling (WJW 2632)] and one sterile 
[39°07'N-70°35'W, 16.viii.1970, Moore (WJW 2910)] — 
contained very small (less than 1 cm long) plants of 



nutans. While 



come 



(Woodward) C. Agardh provided by Taylor (1960, p. 616), 



affinity 



m 



32 



Rhodora [Vol. 77 



Genus Herposiphonia Naegeli, 1846 



Herposiphonia secunda (C. Agardh) Ambronn. Boergesen 
1920:469, Figs. 428-429. Chapman 1963:125, Figs. 130 
a-b. Collins and Hervey 1917:126. Howe 1920:574. 
Taylor 1928:176, pi. 25, Figs. 8-10; 1960:604, pi. 72, 
Figs. 10-11. 

TYPE LOCALITY: LD. 
TYPE: LD. 

distribution: western tropical Atlantic Ocean, Mediter- 
ranean Sea, Adriatic Sea. 

specimens examined: Sargasso Sea: 31°N-69°20'W, 3.iii.l970, 
Volhmann (WJW 2362); 32°09'N-64°58'W, 16.V.1970, Woelkerling 
(WJW 2660); 36°N-70°36'W, 9.xii.l970, Moore (WJW 2935); 37°N- 
70°W, 12.V.1970, Woelkerling (WJW 2630) ; 39°07'N-70°35'W, 16.viii. 
1970, Moore (WJW 2911); 39°30'N-71°W, 6.X.1970, Volkmann (WJW 
2864). 

Male, female and tetrasporangial plants have been found 
on Sargassum fluitans, S. natans, and on Sargassum sp. 

Herposiphonia tenella (C. Agardh) Naegeli. Boergesen 
1918:286, Figs. 287-289; 1920:472, Fig. 430. Chapman 
1963:127, Fig. 133. Collins and Hervey 1917:126. Howe 
1920:573. Taylor 1928:177, pi. 25, Fig. 11; 1960:604, 
pi. 72, Fig. 12. 

type locality: Sicily. 

type: ld. 

distribution : western tropical Atlantic Ocean ; Mediter- 
ranean and Adriatic Seas. 

specimens examined: Sargasso Sea: 33°58.5'N-69°56.5'W, 15.v. 
1970, Woelkerling (WJW 2596); 39°30'N-71°W, 6.X.1970, Volkma'nn 
(WJW 2875). 

The tetrasporangial specimens (WJW 2875) grew epi- 
phytically on Sargassum hystrix; the sterile material was 
found on "Sargassum C." Until specimens attached to S. 
fluitatis and/or S. natans are collected the status of this 
taxon as a permanent component of the Sargasso Sea re- 
mains in doubt. 



1975] Sargasso Sea — Woelkerling 33 



Genus Laurencia Lamouroux, 1813 

Tropical Atlantic American species of Laurencia are in 
need of critical study in light of the recent investigations 
of Saito (1967, 1969). Saito (1967, p. 72-73) recognizes 
5 subgenera of Laurencia based on the presence or absence 
of secondary pit connections in the cortex, shape of cortical 
cells, presence or absence of lenticular thickenings in 
medullary cell walls, and plant form (compressed or cy- 
lindrical). Taylor (1960) does not provide full 
tion on these features, and his keys are based partly on 
features of questionable taxonomic significance, thus mak- 
ing specific identification of taxa d 

Seven collections of Laurencia have been made during 



ma 



fficult 



the present study, but in view of their small size 



mos 



cm 



tion, species affinities have not been determined. One of 
these [34°N-70°W, 30.vi.1970, Volkmann (WJW 2729)] 
belongs to the subgenus Palisadae; the other six [31°N- 
69°29'W, 3.iii.l970, Volkmann (WJW 2361) ; 33°58.5'N- 
69°56.5'W, 15.V.1970, Woelkerling (WJW 2593); 34°N- 
70°W, 6.vii.l970. Volkmann (WJW 2745); 37°N-70°W, 
12.V.1970, Woelkerling (WJW 2636) ; 37°30'N-70°W, 
8.vii.l970, Volkmann (WJW 2710) ; 39°30'N-71°W, 6.x. 
1970, Volkmann (WJW 2874)] belong to the subgenus 
Chondrophycus. The specimens occurred as epiphytes on 
Sargassum nutans and on Sargassum sp. 

Genus Lophosiphonia Falkenberg, 1897 

Lophosiphonia cristata Falkenberg 1901:499, Tab. 9, Figs. 
7-10. Boergesen 1918:297, Figs. 295-298. Chapman 



1964:125, Fig. 129. Hollenberg 1958:68. Taylor 1960: 



606. 



type locality: Scogliera, Puntadel Posilipo, Gulf of 

Napal. 

type: not located. 

distribution: Mediterranean, Bermuda, Bahamas, Ja- 
maica, Virgin Islands, Sargasso Sea. 



34 



Rhodora [Vol. 77 



specimen examined: Sargasso Sea: 31°N, 69°29'W, 3.iii.l970, 
Volkmann (WJW 2363). 

The single collection contained tetrasporangial plants 
epiphytic on Sargassum fluitans. 

Genus Polysiphonia Greville, 1824 

As noted by Taylor (1960, p. 572-3), much taxonomic 
uncertainty exists over tropical American Atlantic species 
of Polysiphonia, and a critical revision of the genus for this 
region is badly needed. Although several sources (includ- 
ing Taylor, 1960) have been consulted during this study, 
taxonomic identifications of Sargasso Sea collections have 
been made mainly from the publications of Hollenberg 
(1968, 1968a). The relationships of Hollenberg's Pacific 
taxa to the species names employed by Taylor (1960) for 
tropical American Atlantic taxa remain uncertain. 
Polysiphonia delicatula Hollenberg 1968:62, Fig. IF. 

TYPE locality: Pokai Bay, Oahu, Hawaii. 
type: us (D1911662). 

distribution: Hawaiian Islands, Tuamotu Archipelago, 
Marshall Islands, Caroline Islands, Sargasso Sea. 

specimens examined: Sargasso Sea: 28°N-70°W, 4.iii.l970, Volk- 
marm (WJW 2423); 33°58'N-69°56.5'W, 15.V.1970, Woelkerling (WJW 
2595); 37°N-70°W, 12.V.1970, Woelkerling (WJW 2631, 3229). 

Female and tetrasporangial plants occurred as epiphytes 
on Sargassum natans and Sargassum sp. 

Polysiphonia lanosa (L.) Tandy. Taylor 1957:341, pi. 56, 

Fig. 4, pi. 57, Figs. 14-15, pi. 59, Fig. 4. Woelkerling 
1972:298. 

Polysiphonia fastigiata auct. non. (Roth) Greville: Col- 
lins, Holden, and Setchell 1895:145; 1907:1444. Far- 
low 1881 : 175. Harvey 1853 :54. 

type locality: unknown. 

TYPE: LINN. 

distribution : North Atlantic Ocean. 

specimens examined: Sargasso Sea: 38°53'N-f>9°39'W, lO.v.1970, 
Woelkerling (WJW 2565). 



1975] Sargasso Sea — Woelkerling 35 



Polysiphonia lanosa occurred as an epiphyte on Asco- 
phyllum nodosum (see Woelkerling, 1972) and is regarded 
as an invader. 

Polysiphonia poko Hollenberg 1968:70, Figs. 3A, 15, 22. 

type locality: North Island, Pacific Ocean. 
type: us (H65-113.1). 

distribution: tropical Pacific Islands (see Hollenberg), 
Sargasso Sea. 

specimens examined: Sargasso Sea: 34°N-70°W, 10.i.l970, Volk- 
mann (WJW 2207); 37°30'N-70°W, 8.vii.l970, Volkmann (WJW 
2703); 39°30'N-71°W, 6.X.1970, Volkmann (WJW 2858). 

Female and tetrasporangial plants occurred as epiphytes 
on Sargassum natans and Sargassum sp. 



SUMMARY 

The epibiotic and pelagic algal flora of the western Sar- 
gasso Sea has been extended to include ten Chlorophyceae, 
25 Phaeophyceae, and 33 Rhodophyceae. Nearly 75% of 
these taxa are newly reported for the Sargasso Sea. Per- 
tinent references and taxonomic and ecological data are 
provided for each taxon, and indication is provided as to 
whether each species is likely to be a permanent component 
of the Sargasso Sea flora or merely a temporary invading 
element. 



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nia. Ibid. 22: 198-207. 

Howe, M. A. 1920. Algae, pp. 553-618. In: N. L. Britton & C. F. 
Millspaugh, The Bahama Flora, vii + 695 pp. New York. 

. & W. D. Hoyt. 1916. Notes on some marine algae 



from the vicinity of Beaufort, North Carolina. Mem. N. Y. 

Bot. Gard. 6: 105-123. 
HOYT, W. D. 1920. Marine algae of Beaufort, N. C. and adjacent 

regions. Bull. Bur. Fish., Wash. 36: 367-556. Pis. I-V. 
Kuckuck, P., & P. Kornmann. 1963. Ectocarpaceen-Studien VIII. 

Einige Arten aus warmen Meeren. Helgolander Wiss. Meere- 

sunters. 8(4): 361-382. 
Kuetzing, F. T. 1843. Phycologia Generalis. (Leipzig). 

. 1849. Species Algarum. (Leipzig). 

. 1854. Tabulae Phycologicae. Vol. 4. xvi + 23 pp. pis. 



1-100. Nordhausen. 



1862. Tabulae Phycologicae. Vol. 12. vi + 30 pp. pis. 



1-100. Nordhausen. 



\Y 



Acta Univ. Lund. (N. F. Avd. 2) 40(2): 1-104. t. 1-32. 
. 1947. Die Phaeophyceen der Schwedischen Westkueste. 



Ibid. 43(4): 1-99. t. 1-18. 



. 1949. Die Chlorophyceen der Schwedischen West- 
kueste. 45(4): 1-79. 

Lanjouw, J., & F. A. Stafleu. 1964. Index Herbariorum. Part I. 
The Herbaria of the World. 5th Ed. Regnum Vegetabile 31. 

Lund, S. 1959. The marine algae of east Greenland. I. Taxo- 
nomical part. Meddr. Groenland 156(1): 1-247. 



1975] Sargasso Sea — Woelkerling 39 



Masaki, T. 1968. Studies on the Melobesioidae of Japan. Mem. 

Fac. Fish. Hokkaido Univ. 16 (%): 1-80. 
Mazoyer, G. 1938. Les Ceramiees de l'Afrique du Nord. Bull. 

Soc. d'Hist. Nat. Afr. Nord. 29: 317-331. 
Nakamura, Y. 1965. Species of the genera Ceramium and Campy- 

laephora, especially those of Northern Japan. Sci. Pap. Inst. 

Alg. Res. Hokkaido Univ. 5(2): 119-180. pis. I-XIV. 
Parr, A. E. 1939. Quantitative observations on the pelagic Sargas- 

sum vegetation of the western Atlantic Ocean. Bull. Bingham 

Oceanogr. Coll. 6(7): 1-94. 
PHAM HoanG-Ho. 1969. Marine algae of South Viet-Nam. 559 

pp. Saigon. 
Prat, H. 1935. Remarques sur la fauna et la flora aux sargasses 

flottantes. Naturaliste Can. 63: 120-129. 
Printz, H. 1964. Die Chaetophoralen der Bennengewasser. Hydro- 

biologia 24(1/3): 1-456. 
Rosanoff, S. 1866. Recherches anatomiques sur les Melobesiees. 

Soc. Imp. Sci. Nat. Math. Cherbourg, Mem. 12: 1-112. 7 pis. 

Rosenvinge, L. K. 1893. Groenlands Havalger. Meddr. Groenland. 

3: 765-981. ph. 1, 2. 
. 1909. The marine alc:ae of Denmark. I. Rhodophyceae 



1. K. Danske Vidensk. Selsk. Skr. (Afd. 7, Raekke) 7(1): 1-151. 
& S. Lund. 1941. The marine algae of Denmark. Vol. 



II. Phaeophyceae. Pt. 1. K. dansk. Vidensk. Selsk. Biol. Skr. 

1(4): 1-79. 

& . 1947. The marine algae of Denmark. 



Vol. II. Phaeophyceae. Pt. 3. K. danske Vidensk. Selsk. Biol. 

Skr. 4(5): 1-99. 
Saito, Y. 1967. Studies on Japanese species of Laiirencia, with 

special reference to their comparative morphology. Mem. Fac. 

Fish. Hokkaido Univ. 15(17): 1-81. pis. 1-18. 
. 1969. The algal genus Laurencia from the Hawaiian 

Islands, the Philippine Islands and adjacent areas. Pac. Sci. 

23: 148-160. 
Sauvageau, C. 1901. Remarques sur les Sphacelariacees. J. Bot. 

Paris 15: 51-167. 

1933. Sur quelques algues pheosporees de Guethary. 



pr 



Bull. Biol. Sta. Arcachon 30(1): 1-128. 29 figs. 
Setchell, W. A. 1931. Hony Kong Seaweeds. II. Hong Kon 

Nat. 2: 237-53. 
SODERSTROM, J. 1963. Studies in Cladophora. Bot. Gothoburgensia 

1: 1-147. Map. 
Taylor, W. R. 1928. The marine algae of Florida with special 
reference to the Dry Tortugas. Pap. Tortugas Lab. 2">: 1-219. 



40 



Rhodora t Vo1 - 77 



1957. Marine algae of the Northeastern coast of North 



Arbor 



. 1960. Marine algae of the Eastern tropical and sub- 



tropical coasts of the Americas. (Ann Arbor). 
van den Hoek, C. 1963. Revision of the European species of Clado- 

phora. Leiden. 
West, G. S. 1908. Some critical green algae. Jour. Linn. Soc. Bot. 

38: 279-289. 
West, W., & G. S. West. 1903. Notes on freshwater algae, III. 

Jour. Bot. Lond. 41: 74-82. 
WlLLE, N. 1900. Algologische Notizen I-VI. Nyt Mag. F. Naturv. 

38: 1-27. 

. 1906. Algologische untersuchungen an der biologischen 

station in Dronthein I-VII. K. Norske Vidensk. Sels. Ske. 

1906(3): 1-39. 
WiNGE, O. 1923. The Sargasso Sea, its boundaries and vegetation. 
Rep. Dan. Exped. 1908-10 to the Mediterranean and adjacent 

seas. 3(Misc) : 1-34. 
Woelkerling, W. J. 1972. Some algal invaders on the northwestern 
fringes of the Sargasso Sea. Rhodora 74: 295-298. 

. 1973. The Audouinella complex (Rhodophyta) in the 



Western Sargasso Sea. Ibid. 75: 78-101. 
Wollaston, E. 1968. Morphology and taxonomy of southern Aus- 
tralian genera of Crouanieae Schmitz (Ceramiaceae, Rhodophyta). 
Aust. Jour. Bot. 16: 217-417. pis. 1-10. 

, 1971. Antithamnion and related genera occurring on 



the Pacific Coast of North America. Syesis 4: 73-92. 
Womersley, H. B. S. 1954. Australian species of Sarfjnssum, sub- 
genus Phyllotrichia. Aust. Jour. Bot. 2: 337-354. 

. 1967. A critical survey of the marine algae of south- 



ern Australia. II. Phaeophyta. Ibid. 15: 189-270. 

& A. Bailey. 1970. Marine algae of the Solomon 



Islands. Phil. Trans. Roy. Soc. Lond., Biol. Sci. 259: 257-352. 



DEPARTMENT OF BOTANY 
UNIVERSITY OF WISCONSIN 
MADISON, WISCONSIN 53706 



SAXIFRAGES ON MOUNT WASHINGTON 

In Rhodora 69: 483-486, three brief articles were pub- 
lished regarding the rediscovery of Saxifraga Aizoon Jacq. 
and Saxifraga cernua L. in Huntington Ravine on Mt. 
Washington. These plants were first discovered by Dr. John 
Churchill in 1939 but were apparently not seen again until 
1967 when Steele, Hodgdon and James Teeri relocated them. 
Although it was not possible to reach the plants because of 
the steepness of the cliff, observations were made through 
field glasses. Saxifraga Aizoon was growing in some abun- 
dance on a nearly flat ledge about twenty feet above a van- 
tage point which was reached without too much difficulty. 
The plants in full bloom with conspicuous typical lime 



unmi 



a ___ _ _„„ the bottom of the cliff. On a 

steep shelf at a distance of forty feet from the vantage 
point, Hodgdon detected a single flower which he believed 
to be Saxifraga cermia, a plant he had become familiar with 
on an Alaskan expedition. Steele, although not familiar 
with the plant was skeptical and unwilling to accept the 
identification. The plant was completely inaccessible so 
there seemed to be no way to resolve the question without 
the aid of rockclimbers. 

Upon learning of the dilemma, Countryman undertook 
to secure the aid of members of Norwich University's 
Mountain Rescue Team. Accordingly, another expedition 
was organized in 1970 consisting of Countryman, Hodgdon, 
Steele and three technical rock climbers from Norwich. 
The vantage point below the Saxifraga Aizoon shelf was 
easily reached but it was then discovered that nearly all 
the soil and vegetation was gone from the ledge, presum- 
ably having been removed by avalanche or flood, a matter 
which emphasized the extreme ecological instability of the 
whole area. The date of this expedition, Aug. 2, was past 
the flowering time of the saxifrages and at first none were 
in evidence. Soon Hodgdon, with the aid of field glasses, 



41 



42 Rhodora [Vol. 77 



was able to locate some plants of S. Aizoon on a ledge above 
the original shelf. Ray Quirk, a rock climbing instructor 
at Norwich, undertook to investigate, assisted by John P. 
Quirk and Jon W. Walsh, both undergraduates at that 
institution. The ledge was very steep and the rocks un- 
stable, but with the use of direct aid in the form of pitons 
and a sling and tension from below, Quirk was eventually 
able to reach the site. Guided by instructions from below 
he was able to obtain small but unmistakable portions of 
both saxifrages, these being the first collections from Mt. 
Washington since 1939. It would have been desirable to 
investigate another ledge but the lateness of the hour pre- 
vented this. 

After this trip, in discussing the 1967 expedition, a 



Hodgd 



•/ 



the time. To settle this point, and because one promising 
shelf still had not been investigated, another expedition was 
organized by Steele in 1971, with a date selected for what 
was believed to be the height of the flowering season of 
both plants. Two rock climbers accompanied the expedition. 
From the vantage point eighty feet up the cliff, S. ccrnua 
was readily observed in flower on a vegetation covered 
shelf below the original S. Aizoon ledge. With the aid of a 
rope and a good belay from Paul Henle, Nathaniel Steele 
was able to reach the slope. He made some careful observa- 
tions and collected one fragment of a plant plus some 

i seven plants of S. cernua in bloom plus 
aves. The slope was steep and unstable 
with many loose rocks. It would not be difficult for all the 
vegetation to be dislodged in a slide. 

Steele (senior) searched the whole area carefully with 
field glasses and eventually located a small clump of S. 
cernua leaves in a small pocket near the original 5. Aizoon 
ledge. On the higher ledge explored in 1970, a considerable 

n was observed in full bloom. The onset 



leaves, 
some i 



amou 



derstorm 



presumably S. cernua was there also. 



1975] Saxifrages — Steele 43 



The narrow gully continues very steeply above this site 
nearly to the head of the ravine, but overhanging rock 
makes it impossible to see into it from the bottom ; thus it 
is possible there are colonies of saxifrage farther up, seeds 
from which might occasionally wash down and colonize 
suitable areas. It would appear this possibility could be 
confirmed onlv by a team of strong rock climbers. 



mi 



rare alpine plants, the authors suggest the following as a 
result of their experiences. High powered field glasses are 



time 



very useful. Careful notes should be taken at the 
what is observed. Drawings and photographs of the locality 
would be most helpful. A camera equipped with a tele- 
photo lens would make an excellent record of the plant. All 
information thus obtained could be put on an herbarium 

sheet. 



seem 



cover new plants in an area that had long been heavily 
botanized. He was fortunate in that he was doing his 
explorations during the short flowering season. His very 
conscientious exploration and mountaineering ability no 



m 



from 



m from 



glasses. 



» manage the last twent 
om a second remain a m 
climbers who have visit 



area. The area is ecologically unstable and there is a slight 
possibility that he collected his plants from a station lower 
down that is now extinct. Because of the instability the 
authors feel that further exploration is inadvisable in that 
it might lead to destruction of the very limited habitat. 

Records for the plants now exist in the herbaria of the 
University of Michigan, the New England Botanical Club, 
the University of New Hampshire, Norwich University and 
the Steele nrivate herbarium in Tamworth, New Hampshire. 



FREDERIC L. STEELE 

THE WHITE MOUNTAIN SCHOOL 

LITTLETON, NEW HAMPSHIRE 03561 



RUDBECKIA AURICULATA (PERDUE) KRAL, 
A SPECIES DISTINCT FROM R. FULGIDA AIT. 



Robert Kral 



During field work toward a flora of Alabama I have come 
across several additional populations of a very tall, smooth- 
ish, rhizomatous Rudbeckia, which was originally described 
by Perdue (1961) from a single population in Covington 
County as var. auriculata of R. fulgida Ait. Admittedly, 
R. fulgida is extremely variable, presenting a broad spec- 
trum of radiate extremes (Perdue treats seven in his 1957 
treatment of the complex) over a broad range of habitats 
and geography. However, this particular Rudbeckia varies 
from the other "fulgidas" so much, and so consistently, that 
to treat it as another variety of R. fulgida would be incon- 
sistent with species concepts in Rudbeckia. Perhaps I can 
best demonstrate this situation by offering descriptions of 
R. fulgida and R. auriculata (R. fulgida var. auriculata 
Perdue), these based on 45 samples of the former and ten 
of the latter. The description of R. fulgida is broadened to 
include the three varieties now known to occur within Ala- 
bama and Tennessee, which are possibly sympatric with 
R. auriculata. 



R. fulgida Ait. var. fulgida. Plate 1(2) ; Plate 2(3). 

R. fulgida Ait., Hort. Kew. 3: 251. 1789. 

R. chrysomela Michx. Fl. Bor. Am. 2: 143. 1803. 

R. truncata Small, Bull. Torr. Bot. Club 25: 478. 1898. 

R. acuminata Boynt. & Beadle, FL S.E.U.S. ed. 1. 1256. 
1903. 

R. foliosa Boynt. & Beadle, Fl. S.E.U.S. ed. 1. 1256. 1903. 
R. tenax Boynt. & Beadle, Fl. S.E.U.S. ed. 1. 1257. 1903. 

Stoloniferous-rhizomatous perennials to 1.3 m. tall, peren- 
nating by one or more slender, stoloniferous offshoots, these 
later thickening and becoming rhizomatous. Rosette leaves 



44 



1975] 



Rudbeckia 



Krai 



15 























' 












■ 

■ 



3 mm 



. 



fa? 

N ill i: ) 

■3 HM y j 




{ 




t * 









v I j 






A 




. . 



, •:■ 



■r ( 




j 



*W**- 



•<? 













«K 
















■ 



L . I 



PLATE I 


















PLATE 1. Pales, akenes and corollas of Rudbeckia. 

1. Rudbeckia auriculata. la. akene; lb. disc corolla; lc. pale, outer 
surface. Drawn from Krai 48579. 

2. Rudbeckia fulgida var. fulgida. 2a. pale, outer surface; 2b. 
akene and disc corolla. Drawn from Krai 33107. 

3. Rudbeckia fulgida var. unibrosa. 3a. akene and disc corolla; 
3b. pale, outer suface. Drawn from Krai 32945. 



46 Rhodora [Vol. 77 



with blades ovate to lanceolate or elliptic, altogether 3-11 
(-25) cm. long-, on petioles shorter than to as long- as the 
blades, the margins ranging from entire to undulate, cre- 
nate, dentate, or serrate, the bases cuneate to attenuate, 
the surfaces glabrous to (usually) appressed-strigose. 
Stems erect, proximally hirsute or strigose or smooth, dis- 
tally strigose, strigillose or smooth, and terete or sulcate- 
costate. Heads few to many, long pedunculate usually from 
the axils of upper stem leaves and forming an open "can- 
delabra-like" cyme. Lower stem leaves similar in outline 
and indumentum to rosette leaves, grading upward to 
shorter, narrower, sessile or even clasping. Phyllaries ob- 
long, slightly to much shorter than the rays, green, ciliolate- 
scabrid, reflexed. Rays short-oblong to elliptic, 0.9-1.7 cm. 
long, spreading or slightly recurved, often strigillose on the 
backs, orange-yellow (frequently drying greenish). Disc 
1.0-1.8 cm. broad, the pales oblanceolate or cuneate, rarely 



acumin 



margi 



surfaces 



cam 



distally a deep reddish-brown becoming proximally tan 
with the 3 nerves, particularly the 2 submarginal laterals 
a deep lustrous reddish-brown, smooth or sparsely strigil- 
lose especially toward the tips. Disc corollas 3.0-3.5 mm. 
long, the tube shorter than the narrowly 
broadly tubular throat, the lobes short-triangular and erect, 
the whole corolla deep purple-brown. Akenes asymmetri- 
cally short-oblong, 2.0-2.5 mm. long, curvate proximally, 
usually 4-angled, dark gray, the pappus a low, slightly un- 
even crown less than 0.5 mm. long or almost obsolete. 

This entity inhabits prairies, swales, open woods, fields, 
and roadsides, on a variety of soils and in dry to quite wet 
situations from New Jersey to Illinois, south to Florida and 
Louisiana. It is definitely the most widesnroad. cwtainlv 



PLATE 2. Habit sketches of Rudbeckia. 

J. Rudbeckia auricula la. 

2. Rudbeckia fulgida var. untbrona. 



• > 



Rudbeckia fulgida var. fulgida. 






PLA TE 2 



48 



Rhodora [Vol. 77 



the most variable taxon of the complex. It and the other 
varieties of R. fulgida all display a fine orange-yellow ligule 
color that distinguishes them well from other Rudbeckias 
in the late summer and early autumn fields. 

R. fulgia Ait. var. umbrosa (Boynt. & Beadle) Cronq., 

Rhodora 47: 400. 1945. Plate 1(3) ; Plate 2(2). 
R.nmbrosa Boynt. & Beadle, Biltmore Bot. Stud. 1: 16. 
1901. 

R. chapmanii Boynt. & Beadle, Biltmore Bot. Stud. 1: 14. 
1901. 

Differs from var. fulgida in being stouter, the stem often 
angular, usually smoother, taller (frequently 1 m. tall), 
with stouter rhizomes. Rosette and lower stem leaves pro- 
portionately large, long-petiolate with ovate or lance-ovate, 
cordate-based blades, these either glabrous or strigose or 
scabrid, and with entire to crenate-dentate margins. Heads 
larger, the rays longer, usually elliptic-oblong, and often 
fully 3 cm. long, the disc often to 2 cm. broad. Pales con- 
sistently ciliate distally save for most inner ones. Akenes 
similar in shape but usually longer, to 3 mm. 

This variety is perhaps the showiest of the Alabama- 
Tennessee fulgidas, in my opinion an excellent horticultural 
subject because of its late and long-blooming habit and its 
full inflorescence. Its best habitats are sunny, moist to 
quite wet swales and seeps from Virginia west to Ohio and 
south to Alabama and Georgia. 

R. fulgida Ait. var. spathulata (Michx.) Perdue, Rhodora 

59:297-298. 1957. 
R. spathulata Michx. Fl. Bor. Am. 2: 144. 1803. 

Most similar to var. fulgida, with rosette and basal stem 
leaves attenuated to the petiole, not cordate. Stems ranging 
from glabrous to variously hirsute or strigose; mid- and 
upper stem leaves ranging from broadly spatulate to ob- 
lanceolate or pandurate, and from glabrous to strigose. 
Heads as small as in var. fulgida with rays short (rarely 
as loner as 2 cm.). 



1975] Rudbeckia — Krai 49 



This variety grows in low, moist to wet, sunny places or 
in low open woods from Virginia west to Tennessee and 
south to Florida and Alabama. In habit, rosette and stem 
leaves, inflorescence, and head it is so similar to extremes 



fulgida var. f 



me 



R. auriculata (Perdue) Krai, stat. nov. Plate 1(1); Plate 

2(1). 
R. fulgida Ait. var. auriculata Perdue, Rhodora 63: 119- 



120. 1961. 



omatous, glabrous or subglabrous perennial 



m 



these becoming stout (to 1 cm. or more thick) by the sec- 
ond season. Rosette and lower stem leaves to 65 cm. long, 
the blades oblong to oblanceolate or ovate-lanceolate, short- 
acuminate to rounded or acute, entire to sinuate, crenate, 
dentate, coarsely serrate or denticulate, on petioles from 
1/3-1/2 the total leaf length, the surfaces smooth to scab- 
rous and usually strigose on the veins, midrib and petiole. 
Stems erect, stiffish, proximally fully 1 cm. thick, subterete, 
multicostate to sulcate. Middle and upper cauline leaves 
sessile, the blades progressively reduced up the stem, ellipti- 
cal to ovate or pandurate, acute, the margins entire to 
dentate, denticulate or coarsely serrate, the bases auricu- 
late-clasping, the surfaces from smooth to sparsely scabrid 
or strumose. Heads many in an open, broadly convex, 
paniculate cyme. Phyllaries oblong-linear, spreading, 
shorter than rays, smooth save for scaberulous margins. 
Pales oblong-cuneate, ca. 6 mm. long, acute, ciliate, the 
backs hirsutulous, proximally tan with maroon mid- and 
sub-marginal nerves, distally a deep reddish-brown. Rays 
bright yellow, oblong-linear, ca. 2 cm. long, spreading or 
spreading-ascending, the backs frequently strigillose. Disc 
1.0-1.7 cm. broad, the corollas purple-brown, 3.0-3.5 mm. 
long, the short tube gradually expanding into the narrowly 
funnelform throat, the lobes short-triangular, erect. Akenes 
oblong-curvate, 4.0-4.5 mm. long, a rich, lustrous reddish- 



50 Rhodora [Vol. 77 



brown, 4 ribbed, the pappus prominently unevenly 4-6- 
toothed, the teeth pale tan, chartaceous, narrowly triangu- 
lar, the longer ones fully 2 mm. long. 

Type: Alabama: covington co.: moist soil along Ala- 
bama Highway 55, 11 mi. south of McKenzie (2 mi. n. of 
Red Level), 24 July 1958, R. E. Perdue 2177. Holotype at 
gh, Isotype at us. 

Rudbeckia auriculata inhabits bogs, swamps, seeps, 
ditches and swales, mostly in full sun, sometimes in partial 
shade at edges of swamp woodlands. So far, most of the 
samples are from the lower Coastal Plain in Alabama, but 
I have found one outlier in the Appalachians, northeast of 
Birmingham in St. Clair County in the broad limestone 
valley between Oak Mountain and Blount Mountain. Oddly 
enough, one of its neighbors there is Leavenworthia exigua 
var. lutea, which appears in patches in spring on low 
outcrops of calcareous rock. Because Alabama is even now 
rather poorly studied for its flora, other populations con- 
necting the montane ones with the Coastal Plain ones will 
probably be discovered. The plants are so tall (some reach- 
ing 3 m. in height!) as to be conspicuous even at a distance, 
the only other Rudbeckia of comparable height in the area 
being R. laciniata. 

Rudbeckia auriculata, to summarize, differs from R. ful- 
(jida in the following ways : 

1. The plants are usually at least twice as tall, arising 
from much stouter rhizomes, and are prevalently smooth- 
stemmed. 

2. The foliage is, overall, larger, with cauline leaves 
prevalently panduriform, auriculate-clasping; this char- 
acter is stressed by Perdue (I.e.) in his varietal distinction. 
The phyllaries are hairy on the backs with spreading or 
ascending hairs. 

8. The rays are a bright yellow, not an orange-yellow as 
in the fulgidas of the southeastern U.S. It is to be suspected 
that their pigment chemistry is quite different. 



1975] Rudbeckia — Krai 51 



4. The disc corollas are shorter than the akenes. In Rud- 
beckia fulgida the reverse is true, i.e., the akenes are 
shorter than the disc corollas. 

5. The akenes are narrower in outline and longer than 
are those of Rudbeckia fulgida, and are a lustrous red- 
brown rather than gray-tinted. 

6. The pappus is prominent, produced upward into nar- 
rowly triangular scales, these longer than is true for any 
other southeastern Rudbeckia. In R. fulgida the pappus is 
usually a low, uneven crown or is nearly absent. 

Thus, when one considers the many striking character 
differences, this is not only a species distinct from Rud- 
beckia fulgida, but indeed might well comprise a part of 
another section of the genus. Only a revisional attempt 
would disclose this fact, but the taxa nearest to R. auricu- 
lata in pappus character and in some respects of head char- 
acteristics are. in southeastern U.S.. R. nitida and R. mohrii. 



A cytological comparison of some southeastern Rud- 
beckias is to be initiated during the summer of 1974, for 
there is sufficient phenotypic difference to assume a con- 
siderable chromosomal difference, at least from R. fulgida 
and its varieties. 



The following are some recently collected localities for 
populations of Rudbeckia auriculata: 



Alabama: barbour co.: swampy ditch beside low oak forest, Lugo, 
12 Sept. 1968, R. Krai with K. E. Blum 33300. Covington CO.: tall 
plants of peaty swale by Ala. 55, just n. of Red Level, 30 Au#. 1970, 
Krai 40841 ; sandy silt of creek bottoms by Ala. 54, 7.8 mi. n.e. 
Flerala, plants to 8' tall, 24 Jul. 1968, Krai 31970. Crenshaw CO.: 
n. side Luver^ne, sandy creek bottom by U.S. 231, plants to 3 m., 
16 Aug 1 . 19(58, Krai 32421. GENEVA Co.: sandy peat of longleaf pine- 
land ditch duo e. of Samson, just outside limits by Ala. 52, 3 Sept. 
1969, Krai 36837; 5 mi. s. of Samson along- Ala. 187, moist ditch, 
tfrown up waste area, roadsides, 18 Au^. 1970, H. D. Moore 640. 
pike CO.: sandy shores of Pike Co. public lake, s. of Troy, 11 Sept. 
1968, Krai 33174. ST. CLAIR CO.: ca. 4 mi. n. Leeds, abundant clone 
in wet cleared area of pasture, 27 Sept. 1972, Krai 48579. 



52 Rhodora [Vol. 77 



Field work toward this paper was supported largely by a 
research grant to the author from the National Science 
Foundation (NSF GB-6688X). 



References 

Perdue, R. W., Jr. 1957. Synopsis of Rudbeckia subgenus Rud- 



beckia. Rhodora 59: 293-299. 



63: 119-120. 



fulgida. Rhodora 



Small, J. K. 1903. Flora of the Southeastern United States. 1394 

pp. New York, published by the author. 
. 1933. Manual of the Southeastern Flora. 1554 pp. Re- 



print, Univ. of North Carolina Press, Chapel Hill. 



BIOLOGY DEPARTMENT 
VANDERBILT UNIVERSITY 
NASHVILLE, TENNESSEE 37235 



APPARENT ECOTYPIC DIFFERENCES 

IN THE WATER RELATIONS OF 
SOME NORTHERN BOG ERICACEAE 1 



Peter J. Marchand- 



The apparent paradox of xeromorphy in northern bog 
plants has become a popular subject again, this time how- 
ever, with a new interpretation. The development of 
sclerophyllous leaf tissue (increased cutinization and lig- 
nification, increased fiber/protein ratio) in plants of both 
wet and dry habitats has recently been linked with phos- 
phorous deficient soils (Beadle, 1966; Loveless, 1962), a 
feature that both deserts and northern bogs have in com- 
mon. In the light of this discovery, leaf sclerophylly and 
the evergreen habit often associated with it are now being 
interpreted as reflecting a specialized metabolism tolerant 
of low nutrient levels, though the physiology of this adap- 
tation is not yet fully understood (Small, 1972a, 1972b; 
Beadle, 1968, 1966; Monk, 1966; Loveless, 1962). 

From a morphological viewpoint much of the northern 
bog flora seems particularly well adapted for the conserva- 
tion of water. Narrow, leathery, and heavily cutinized 
leaves with revolute margins and vertical orientation, char- 
acters typical of many Ericaceae, are perfect water con- 
serving adaptations. Yet these features cannot be explained 
on the basis of plant-water relations in the northern sphag- 
num bog. Early hypotheses relating these xeromorphic 
characters to a condition of "physiological drought," for 
reasons mostly associated with the anaerobic bog substrate, 
were dismissed long ago by Caughey (1945) for lack of ex- 
perimental confirmation. More recent in situ determinations 
of the water status of bog plants by Small (1972a) and 



'Contribution No. 2, The Center for Northern Studies. 
-Present address: The Center for Northern Studies, Wolcott, Ver- 
mont 05680. 



53 



54 Rhodora [Vol. 77 



Marchand (1972) have likewise failed to turn up any evi- 



environment. While 



middav xvlem 



found to be quite low ( - 15 to —20 bars) , stomatal response 
appears to be unaffected in this range (Small, 1972a; Mar- 
chand, 1972). Assuming that stomatal functioning is an 
adequate measure of plant sensitivity to its environment, 
there seems enough justification indeed to abandon our 
ideas of water stress in bogs. 

This is not to suggest, however, that the subject of water 
relations of bog plants is closed. To the contrary, the acid 



some 



problems 



(1972a), 



as well as that of Marchand (1972), invite further investi- 
gation into possible differences in the water metabolism of 
bog plants and their counterparts on well drained sites. 
Specifically, the following questions may be asked: Why 
are xylem potentials of small bog shrubs so low, sometimes 
exceeding -20 bars, when soil water supply in this habitat 
is seemingly unlimited? In an environment where soil- 
water supply is more restricted, how low a water potential 
might these species tolerate, before stomatal closure is 
induced? Do the observed water potentials represent an 
optimum "operational" level for the species in question or 
is the level induced by 

habitat? Presented here are some field observations which 

suggest possible answers to these questions. 



some 



METHODS 



H 



num 



by several members of the family Ericaceae along with 
Picea mariana (Mill.) BSP. and Eriophorum virginicum L. 
Because the bog is situated in a kettle amidst glacial de- 
posits, however, the surrounding upland area contrasts 
abruptly in both edaphic character and floristic composition. 
Around the perimeter of the bog, the well-drained till sup- 



1975] Bog Ericaceae — Marchand 55 



--■# - 



ports a "mesic" pine-oak community. Both the bog and 
adjacent upland sites, however, support well established 
populations of Vaccinium corymbosum L. and Gaylussacia 
baccata (Wang.) K. Koch, thus affording an opportunity 
to study simultaneously the water relations of these species 
under similar atmospheric conditions but under markedly 
different edaphic conditions. In Wolcott, Vermont, a some- 
what similar situation exists in which the water status of 
Ledum qroenUindicum Oeder and Kalmia angnsti folia L. 



com 



same 



At each site, xylem potentials were measured using a 
portable pressure bomb (PMS Instruments Co.) as de- 
scribed by Waring and Cleary (1967). The pressure bomb 
is designed to measure the amount of tension in the xylem 
water column resulting from transpirational "pull" against 
flow resistances in the soil-plant continuum. This xylem 
tension is related to the free energy of the water in the 
plant, the water potential. When the stem of a transpiring 
plant is cut, the cohesion of the water column is broken 
and the tension is relaxed. The water column then quickly 
recedes from the cut surface. By placing the branchlet into 
a sealed chamber with only the cut end protruding, and 
applying pressure, the water column can be forced back to 
the cut surface, thus re-establishing the initial condition. 
The amount of pressure required to do this is equal in 



xylem 



of the plant. 



cm 



and bearing several leaves were used for the water poten- 
tial determinations. The sample number ranged from just 
three stems per species when variation was low, as during 
the early morning hours, to six when the variation within 
species was higher. At the time of sampling, soil-water 
potentials were determined using a tensiometer (Soilmois- 
ture Equipment Corp. No. 2900) and wet- and dry-bulb 
temperatures were recorded with a sling psychrometer. 



56 Rhodora [Vol. 77 



Since pressure bomb measurements by themselves are 
difficult to interpret, stomatal response to the observed 
water potentials was measured with a diffusion porometer. 
This instrument is essentially an electric hygrometer whose 
resistance varies inversely with humidity. In use, a small 
plexiglass chamber containing a hygroscopic element is 
clamped onto a leaf and, in effect, the leaf is allowed to 
transpire under prescribed conditions. As water vapor 
diffuses into the initially dry chamber, the time rate of 
humidity increase over a narrow range is noted. From this, 
the diffusion resistance of the leaf, mostly a function of 
stomatal aperture, is calculated based on a calibration 
curve for known resistances. 



om 



design of Kanemasu et al. (1969). In order to increase 
instrument sensitivity for use on very small leaves the 
chamber volume was reduced slightly, a higher sensitivity 
element (Hygrodynamics No. 4-4816K) was used, and the 
cup aperture was reduced to 6.35 mm. diameter. With 
these modifications, the time required for attachment and 
measurement on the leaf was usually of the order of 5 to 15 
seconds when the stomates were fully open. In calibration, 



Slatyer (1971a, 1971b) were followed closely. 



M 



RESULTS 



The data from the different sites are compared by using 
the model of Elfving et al. (1972) where transpiration is 
described by the ratio of vapor pressure deficit (VPD) to 
stomatal diffusion resistance (r s ). A plot of xylem water 
potential against VPD/r s for the species studied here indi- 
cates that for any given transpiration rate, under non-stress 

potentials are lower (more negative) in 
the bog plants than in their upland counterparts (Fig. 1). 

The rapid development of a very steep water potential 
gradient in the bog plants was observed early in the day. At 
the Lee Hill Bog, where soil water potential was zero bars, 



em 



1975] 



Bog Ericaceae — Marchand 



57 






\> 








O 





t/> 



CVI 




T 



T 



T 



T 



I 



T 



T 



SJP 




X 





/ 



T 



T 



T 



T 



~r 



T 



I 



I 



A 



Figure 1. Xylem water potential (v^x) as a function of estimated 
transpiration rate (VPD/rs) for bog* and upland ecotypes under non- 
stress conditions. Symbols: O L. groenlandicum ; D K. angustif olia ; 
■ G. baccata; V corymbosum. 



58 



Rhodora [Vol. 77 



from 



ing the first hours after sunrise by a heavy dew formation 
on the leaves. As soon as the dew evaporated and trans- 
piration began, a very steep gradient was established with 
xylem water potential decreasing from —1 to —15 bars 
within one hour and further decreasing to -22 bars in the 
next four hours (Fig. 2). The leaf stomates remained open 
(r s = 1.1 sec/cm.) throughout the day however. In the 



water notential was —0.3 



from 



xylem water potentials decreasing gradually, eventually to 
be checked by an increase in stomatal resistance to an 
average of 4.8 sec/cm. by mid-morning and 8.3 sec. /cm. 

by mid-afternoon (Fig. 2). 

While a soil-water potential of —0.3 bars is not usually 
considered very dry, the difference in soil-water availability 
between the above two sites is best indicated by the slow 
rate of recovery or increase of xylem potential in the up- 
land population as compared to that of the bog population, 
beginning with the approach of sunset and continuing to 
daybreak (Fig. 2). In spite of such differences in soil- 
water availability, stomatal behavior, and transpiration 
rate (an increase in r s from 1.1 sec/cm. to 8.3 sec/cm. is 
enough to cause a five-fold decrease in the transpiration 
rate), the mid-day depression of water potential in both 
populations was similar: —19.4 ± 2.8 bars in the upland 
population and —22.0 ± 2.7 bars in the bog population 
(Fig. 2). 



DISCUSSION 

When the rate of movement of water through the plant 
falls behind the rate of transpiration loss at the leaf sur- 
faces, due either to internal resistances to flow or an in- 
sufficient soil-water supply, the result is a decrease in plant 
water potential. If there were no internal resistances in 
the plant, then in any situation where soil-water is non- 
limiting, the plant water potential would always be very 
high (near zero). Conversely, where soil-water is freely 



1975] 



Bog Ericaceae — Marchand 



59 



if) 





o 




<D 





CO 



^T 



C\J 




O 




00 



CD 



CD 



CD 



^r 



C\J 




o 



LO 



o 



in 



i 





i 



i 



o 

i 



if) 

i 



(sueq) x a 



Figure 2. Diurnal trend of xylem water potential (^x) in Gay- 
hissacia baccata, Lee Hill Bog and adjacent south-facing- hillside. 
Stomatal diffusion resistance (r s ) is given in sec/cm. Vertical bars 
represent standard deviations. 



60 



Rhodora [Vol. 77 



available and xylem water potentials become appreciably 
low (more negative), as in the bog community, then in- 
ternal resistances are indicated. 

For steady-state flow through the plant, the transpiration 
flux is considered to be directly proportional to the water 
potential gradient and inversely proportional to internal 

Therefore, relating the measured xylem 
to the transDiration rate as estimated by 



flow resistances. 



the ratio VPD/r s (for non-stress conditions) indicates that 
flow resistances within the bog plants may be higher than 
in the upland ecotypes. The rapid decrease of xylem poten- 
tial with the commencement of transpiration in the bog 
plants gives some measure of the magnitude of the driving 
force (water potential gradient) needed to move water 
through the plant in the presence of high internal resist- 
ance. 

Assuming that there is no resistance to water movement 
in the saturated peat soil, the resistances indicated for these 
bog plants may be considered to lie between the root-soil 
interface and the stem xylem tissue just beneath the leaf 
layer (the point at which the xylem water potential was 
measured) . It is believed that poor soil aeration, a condition 
typical of bogs (see, for example, Armstrong and Boatman, 
1967), inhibits root absorption, but is this alone enough to 
account for the resistance indicated here? Studies of Mack- 
Ion and Weatherley (1965) suggest that root resistance 
plays little part in establishing leaf water potentials lower 
than — 5 bars. 

Flow resistances in the xylem, on the other hand, are 
generally considered to be the lowest in the soil-plant- 
atmosphere continuum. Increased resistance in the xylem 
may possibly result from decreased diameters of conducting 
elements or through the blocking of these by sclerids or 
tyloses, creating in effect a more tortuous translocation 
pathway in bog plants. Preliminary microscopic examina- 
tion of the xylem tissue, however, showed vessel elements 
to be on the order of 15 to 25 microns in diameter, essen- 
tially the same as in the upland ecotypes and not small 



1975] Bog Ericaceae — Marchand 61 

enough to increase flow resistance, and revealed no obstruc- 
tions in the vessels (Terrell Comstock, unpublished data) . 
The source of the high internal flow resistances in bog 
plants, if indeed they exist, thus escapes recognition for 



moment 
The values for VPD in this 



om 



the wet-bulb depression only (i.e., atmospheric VPD) 
which presupposes that the leaf and air were in thermal 

If leaf temperatures were elevated above air 



mm 



temperature, then the calculated values for VPD/r s would 
be low. Presumably however, with all plants fully exposed, 



same 



Moreov 



mu 



assum 



their roots is not saturated, then the relative differences in 
water potential levels seen here take on even greater sig- 
nificance with respect to indicated internal differences. 

Of considerable interest in these data is the fact that the 
midday xylem potentials in the two Gayhissacia populations 
remained relatively close. This suggests the possibility of 
a closely regulated optimum water potential level for the 
species, indicating also very sensitive stomatal control, with 
the stomates of the upland plants closing before any de- 
tectable decrease in xylem water potential. This same 
phenomenon has been observed in yellow birch (C. A. 
Federer, personal communication) and is somewhat sug- 
gestive of the kind of direct response of stomates to evapo- 
rative conditions of the atmosphere, independent of leaf 
water potentials but conditioned by soil water supply, as 
reported by Schulze et al. (1972). 

As for the question of xeromorphy in bog plants, leaf 
sclerophylly appears of advantage only insofar as it may 
be related to the evergreen habit, possibly preventing win- 
ter desiccation when water uptake is impaired. There is not 
enough evidence yet, however, to conclude that the water 
metabolism of bog plants is entirely similar to that of their 
upland counterparts. While Small (1972a) demonstrated 



62 Rhodora [Vol. 77 



some similarity in the water 



bog and nearby mesic habitat, his mesic site was described 
as including a marsh and field adjacent to the bog and 
many of the species studied at that site, such as Ilex verti- 
ciUata (L.) Gray, Nemopanthus mucronata (L.) Trel., and 
Alnus rugosa (DuRoi) Spreng. are typically confined to 
wetland areas. Lacking any data to the contrary, this sug- 
gests that the two environments may not have differed 
significantly in terms of soil-water availability. The data 
presented here indicate that dissimilarities in water metab- 
olism between bog and upland ecotypes may indeed exist in 

relation to soil-water availability or other edaphic differ- 
ences. 



ACKNOWLEDGEMENTS 



This study was supported in part by funds provided by 
the Office of Water Resources Research, U.S. Department 
of the Interior, by a Grant-in-Aid of Research from the 



for Northern Studies, Wolcott, Vermont 



Center 



Literature Cited 



Armstrong. W 



1967. Some field observations 



relating the growth of bog plants to conditions of soil aeration. 

Jour. Ecol. 55: 101-110. 

Beadle, N. C. W. 1968. Some aspects of the ecology and physiology 
of Australian xeromorphic plants. Aust. Jour. Sci. 30: 348-355. 



1966. Soil phosphate and its role in molding segments 
of the Australian flora and vegetation, with special reference to 
xeromorphy and sclerophylly. Ecology 47: 992-1007. 
Caiigiiey, M. G. 1945. Water relations of pocosin or bog shrubs. 
PI. Physiol. 20: 671-689. 

Elfving, P. C, M. R. Kaufmann, & A. E. Hall. 1972. Inter- 
preting leaf water potential measurements with a model of the 
soil-plant-atmosphere continuum. Physiol. Plant. 27: 161-168. 

Kanemasu, E., G. Thurtell, & C. B. Tanner. 1969. Design, cali- 
bration and field use of a stomatal diffusion porometer. PI. 
Physiol. 44: 881-885. 

Loveless, A. R. 1962. Further evidence to support a nutritional 

interpretation of sclerophylly. Ann. Bot. 26: 551-561. 



1975] Bog Ericaceae — Marchand 63 



Macklon, A., & P. E. Weatherley. 1965. Controlled environment 
studies of the nature and origins of water deficits in plants. 

New Phytol. 64: 414. 
Marchand, P. J. 1972. Stomatal resistance and transpiration po- 
tential of bog plants. Unpublished Master of Science Thesis, 

Univ. N.H. 
Monk, C. D. 1966. An ecological significance of evergreenness. 

Ecology 47: 504-505. 
Morrow, P. A., & R. 0. Slatyer. 1971a. Leaf temperature effects 
on measurements of diffusive resistance to water vapor transfer. 

PI. Physiol. 47: 559-561. 
& . 1971b. Leaf resistance measurements 



with diffusion porometers: Precautions in calibration and use. 

Agr. Meteorol. 8: 223-233. 
Schulze, E. D., O. L. Lange, U. Buschbom, L. Kappen, & M. 

Evenari. 1972. Stomatal responses to changes in humidity in 
plants growing; in the desert. Planta 108: 259-270. 

Small, E. 1972a. Water relations of plants in raised sphagnum 
peat bogs. Ecology 53: 726-728. 

. 1972b. Ecological significance of four critical ele- 
ments in plants of raised sphagnum peat bogs. Ecology 53: 498- 

503. 
Waring, R. H., & B. D. Cleary. 1967. Plant moisture stress: 

evaluation by pressure bomb. Science 155: 1248-1254. 



DEPARTMENT OF BOTANY AND PLANT PATHOLOGY 
UNIVERSITY OF NEW HAMPSHIRE 
DURHAM, NEW HAMPSHIRE 03824 



THE DECIDUOUS MAGNOLIAS 

OF WEST FLORIDA 



Ronald F. Miller 



The deciduous magnolias are perhaps the most distinctive 
relicts of the old Arcto-Tertiary forest. Pleistocene glacia- 
tion apparently eliminated these once-widespread plants 
everywhere from the temperate zone except in the eastern 
United States and in an arc stretching from Sikkim and 
Nepal to the Japanese Islands. Abundant rainfall, moderate 
temperatures, and rich mixed hardwoods seem essential for 
their greatest development in such regions as the southern 
Appalachians and the Cumberland^ in this country. There 
Magviolia acuminata and M. fraseri contribute a minor 
though noteworthy element to the mixed mesophytic forests, 
growing alongside tuliptrees, hemlocks, lindens, buckeyes, 
and other characteristic species. In the lower valleys, 
especially along watercourses, M. tripetala and (in the 
Cumberlands) M. macrophylla display their huge leaves. 
Even the casual observer soon learns to associate the 
deciduous magnolias with our mountain forests in their 
cool, well-watered lushness. It may come then as a surprise 
to discover that each of these four eastern deciduous mag- 
nolias, either in itself or in a closely related species, can 
be found amid the predominantly piney woodlands of the 
Florida Panhandle. 

This is not to suggest that these species can be found on 
the Coastal Plain only in West Florida, but that their re- 
markable collocation and the mode of their survival there 
in a presumably alien environment are of special taxonomic 
and ecological interest. This region differs from the rest of 
the lower Coastal Plain stretching from Virginia to eastern 
Texas mainly in its deeply incised terrain. The pine-covered 
uplands and the evergreen bays characterizing this zone 
are interrupted here and there by bluffs, steep-sided water- 
courses, and sharply rolling country, all of which provide 



64 



1975] Magnolias — Miller 65 



a foothold for a strongly deciduous and often surprisingly 
Appalachian plant assembly. 1 On these slopes, especially 



moi 



genera such as Fagus, Halesia, Oxydendrum, and Tilia mix 
with such subtropical species as Magnolia grand i flora and 
Que reus laurifoliaf some of the riverbluffs supported 
American chestnut until quite recently (Elias, 1971). : The 
understory will often prove to be a thick tangle of the 
Appalachian Kalmia latifolia and the coastal bay-plant 



floridanum 



where 



com 



such as Trillium and Sanguinaria put in an appearance. 



mesic 



cam 



These anomalous patches of forest suggest a time, un- 
doubtedly during a glacial epoch, when a deciduous forest 
covered much more of the area. Phytogeographers have 
inn a- rmtpd the more northerlv elements in the ravine flora 



'See Thorne (1049) for ;i discussion of Appalachian, species in the 
ravines of Southwest Georgia. E. L. Braun (1950) mentions the 
presence of a more deciduous element in the incised Tunica Hills 
section of eastern Louisiana. 

-There seems to he some confusion about the proper nomenclature 
for the common "laurel oak" of the coastal zone. Kurz and Godfrey 
(1902) argue that this plant should be called Q. hemisphaerica 
Bartram. Tn this paper I have chosen to follow the conventional 

ascription of Preston (1961). 

^Old-timers in Okaloosa and northern Escambia Counties recall the 
chestnut tree quite well. Unfortunately, the crowns do not seem to 
sprout as well as they do further north, and the frequency of chink- 
apins in the area keeps the blight pathogen in plentiful supply. 

'Walter's pine (Pinus glabra) is also always present in these 
deciduous forests, but it is net a good indicator since it is also an 
element of what may be called the hammock association, a mixture 
dominated by this pine, Magnolia grandiflora, and the evergreen 
oaks Quercus virginiana and Quercus laurifolicu This climax associa- 
tion will eventually take over dry pine flats and ridges if the leaf 
litter is allowed to build up and if the woods are protected from fire 
and lumbering. The hammock association intergrades with the de- 
ciduous forests on more gradual slopes, especially where the soil is 
quite sandy. 



66 Rhodora [Vol. 77 



of the famous Apalachicola River bluffs in Gadsden and 
Liberty counties, with their endemic Torreya and Taxus 
and such Appalachian species as Cornus altemifolia. A 



endemic such 



mam 

more 



(see James, 1961). Apparently only steep and (often) 
north-facing terrain offered sufficient protection from the 
hot winds and intense insolation that favored the develop- 
ment of evergreen oaks and pines instead of deciduous 
woodlands. Only in these protected enclaves do the decidu- 
ous magnolias maintain their precarious Floridian existence. 
In a few years clear-cut silviculture and residential 
sprawl may make fieldnotes on the adaptations of these 
plants to an alien environment unobtainable. The follow- 
ing observations on various species are offered for whatever 



m 



Magnolia ashei Weatherby 



American m 



m 



been open to dispute since its description by C. A. Weath- 
erby in 1926. Weatherby differentiates it from Magnolia 
macrophylla by rather comparative traits: a little less hair 
on the undersides of the leaves, smaller flowers and leaves, 
the smaller size of the plants. Only its rather narrow in- 
fructescence, as opposed to the strikingly globular one of 
M. macrophylla, offers any certainty to the identification 

specimens (Kurz & Godfrey, 1962). I am 
personally very familiar with both plants, yet am not at 
all sure I would be able to distinguish a vigorous leaf or 
large flower of M. ashei from one of ordinary M. macro- 
pin// la. Even the variations in the purple blotches at the 
center of the flowers — something rather difficult to observe 
in herbarium specimens — seem to run through identical 
ranges in the two plants. 

The two plants are quite separable in the field, however. 
Magnolia macrophylla is essentially a tree; it competes for 
space in the canopy in much the same wav a hir.korv or n 



1975] Magnolias — Miller 67 



sweetgum will do, though these taller-growing trees may 
eventually overtop it. The young plants are therefore re- 
markably upright, and blossoming occurs only on mature 
shoots high above the ground. The collector who wishes 
to examine a M. macrophylla flower must locate either an 
isolated horticultural .specimen or one released by clearing 
or roadbuilding. Magnolia ashei on the other hand behaves 
as a large, coarse shrub that seldom climbs out of the 
understory. It competes for light in the manner of the 
deciduous azaleas, by leaning toward holes in the canopy 
and by taking on a loose-jointed, often horizontal growth 
form. On rare occasions an individual specimen will become 
what might be termed a tree, but that plant will hardly 
ever exhibit the clean upward sweep of the average M. 
macrophylla. Though the plant's crowns sprout vigorously, 
individual shoots of M. ashei, from the evidence of all the 
dead wood, seem to be quite short-lived, and the species 
apparently is incapable in the wild of that sustained devel- 
opment on a single shoot necessary for competition in the 
upper layers of the forest. Horticultural specimens of M. 
macrophylla grow continuously from a single bole if undis- 
turbed; M. ashei specimens can hardly be kept from cop- 
picing. 

Magnolia ashei seems to have adapted to its existence as 
a shrub by assuming a tolerance for shade and by blooming 
in the understory. The plant is remarkably free-flowering 
at small sizes; in the wild I have observed small shoots (ca. 
1 meter) capped by a blossom while standing in the dense 
shade of Quercus lauri folia. In cultivation, the plant be- 
comes strikingly floriferous. One in my care has set blooms 
on coppice shoots less than 10 cm. tall, and another has 
borne five blossoms on a single branched shoot not one 
meter tall or half a meter across. Such behavior in M. 
macrophylla is, to the best of my knowledge, unheard-of. 

The present-day ranges of these two plants preclude 
genetic exchange. In Alabama I have never observed 
Magnolia macrophylla south of the rolling limestone hills 
stretching from northern Mobile County to Monroeville 



68 



Rhodora [Vol. 77 



and then eastward. In Mississippi and Louisiana the plant 
goes further south into areas underlain by more recent 
Tertiary formations, but along the Alabama-Florida border 
only a few north-south river valleys and bluffs interrupt the 
sterile Citronelle (late Pliocene or early Pleistocene) and 
later Pleistocene deposits blanketing the terrain. Neither 
taxon seems to have invaded the few upland hardwood areas 
in this region, possibly because both prefer rather more 
calcareous soils. I have never seen M. ashei north of the 
line traced by U. S. Highway 90. 

The center of distribution of Magnolia ashei seems to be 
the Knox Hill region in eastern Walton County. In this 
strongly rolling area the sterile overburden is replaced by 
a dark, rich, water-retentive clay or loam derived from 
distinctly calcareous marl. 5 The plant is actually common 
on a few square miles surrounding Knox Hill. To the east 
it can be found on hills and along watercourses near Ver- 
non, Washington County (geologically a rather similar 
area) ; on limestone hills along Econfina Creek in northern 
Bay County; on high slopes along the east bank of the 
Ochlockonee River near Smith Creek, Wakulla County; and 
on the Apalachicola bluffs between Bristol and Chattahoo- 
chee. The stations a few miles south of Chattahoochee bring 
M. ashei closest to M. macrophytta : patent M. macrophylla 
may be found in the ravines south of Ft. Gaines, Georgia, 
some 50 miles to the north. As best I can discover, the 
southern border of Clay County, Georgia, marks the south- 
ernmost extension of M. macrophylla, so intergradation 
seems highly unlikely. 11 It may be of interest that two other 
separable but closely related taxa, Rhododendron minus 
and its Florida form, R. minus var. chapmanii, reach their 



3 The region contains rich collecting spots for Miocene shells of the 
Alum Bluff series. See Cook and Mosson (1929). Harper (1914) 
describes the soil and vegetation of this area* 

6 At least in Georgia. I am much less familiar with the southeastern 
corner of Alabama, so intermediates could conceivably be found in 
Htnry or Houston counties. If they are there, however, they are 
elusive. 



1^75] Magnolias — Miller 69 



southernmost and northernmost stations within less 



M 



miles of the comparable stations for M. macrophylla and 



Hill region. M 



within 



Eglin Air Force Reservation. The westernmost stat 

just off State Highway 87, about 20 miles east of Pensacola. 
It was from this region that W. W. Ashe obtained the 
series upon which Weatherby based his description of the 
species. Oddly enough, the soil on these slopes is sandy 
and apparently sterile, quite unlike the heavy soil of sta- 
tions to the east. The deciduous element of the attendant 
vegetation is also attenuated, with Fagus notably absent, 
though Ostrya, Oxydendrum, and Stewartia are almost 
always present. Gordonia lasianthus is quite frequent 
along these streams. 7 

Magnolia ashei is thus probably a relict that through iso- 
lation has adapted to life as a shrub rather than a tree. 
The plant inhabits mixed evergreen and deciduous forests, 
and indeed the dense canopy offered by the evergreen mag- 
nolia and the evergreen oaks may have precluded competi- 
tion in the manner open to M. macrophylla in regions where 
these evergreen hardwoods are not so common. Except in a 
few square miles the plant is remarkably rare; a compar- 
ably vulnerable, unusual, and attractive bird or mammal 
would surely have been the occasion for dozens of pilgrim- 
ages and popular articles. 

7 The presence of Magnolia ashei in East Texas has been reported. 
Several years ago I observed a big-leafed magnolia in the yard of a 
gentleman living at "Devil's Pocket," a rather flat, swampy region 
in southern Newton County. He said this transplanted specimen was 
all that remained of an older plant that once stood in his pasture 
on a hammock. The plant has since died, so I have been unable 
to re-examine the specimen in the light of my field experience with 
Florida M. ashei. As I recall, the plant bore reduced leaves, but the 
form was definitely upright in a manner I later learned to associate 

1 ft ■ ^ ■ m * m _ 



Mag 



and conferring have fa 
county or in Texas. Magn 



infructescences. Several 



noted, is fairly common in adiacent western 



70 



Rhodora [Vol. 77 



Magnolia acuminata L. 

Magnolia acuminata is a more decidedly Appalachian 
plant than M. macrophylla, yet it occurs in West Florida, 
perhaps in the form of M. acuminata var. subcordata. The 
distinction between the typical variety and var. subcordata 
seems more plausible to the reader of taxonomic keys than 
to the observer in the field, since great variations in hairi- 
ness and flower color may occur within a given locality 
(see Hardin, 1954). In general, the West Florida plants 
bear greenish-yellow flowers intermediate between the 
showy canary-yellow blooms of some plants in piedmont 
Georgia and the greenish forms prevalent in the higher 
mountains. Although the hairiest leaves I have ever seen 
on any cucumber-tree grew on a Florida specimen, closeby 
trees bore leaves indistinguishable from those on central 
Mississippi and northern Alabama plants. As with many 
other species, hairiness seems to increase gradually though 
irregularly southward, as water loss from the leaves be- 
comes more and more of a threat to survival. I wonder if 
the cucumber-trees would be split at all if it were not for 
the historical accident that a singularly yellow form existed 
as a horticultural curiosity for almost a century until the 
piedmont Georgia plants were relocated by Berckmans in 
1913 (Sargent, 1933). 

In Florida Magnolia acuminata is to be found only in the 
Knox Hill region mentioned in the previous discussion. It 
is much less frequent there than M. a&hei, however, growing 
only in a narrow band marking the transition between 
beech and hickory woods along a few ravines and slopes in 
the richest areas. There it becomes a fairly large tree. 
Kurz and Godfrey (1962) report that a search of the area 
located only six trees. Though I have located several 
hundred, distribution is sporadic and the trees are easily 

missed without a meticulous search in rather difficult 
country. 

The Citronelle deposits mentioned earlier separate these 
Florida plants from the closest Alabama cucumber-trees, 



1975] Magnolias — Miller 71 



a large population inhabiting the Sepulga River valley to 
its mouth in upper Escambia County, Alabama. The mouth 
of the Sepulga marks the southernmost outcropping of 
Oligocene limestone in that part of the state, so calcareous 
soil aerain seems to be the key to distribution. The 



lime 



Marianna 



see apparently does not support cucumber-trees, but then 



cum 



sent in immediately adjacent Alabama and southwestern 
Georgia. Southwestern Alabama and southern Mississippi, 



ucumber-trees. Possi- 



anomaly elude me 



Genetic isolation seems to have resulted in no genetic 
drift: trees from western Florida and southern Alabama 
occupy virtually identical sites and, from every appearance, 
are indistinguishable. Magnolia acuminata seems to be an 
intermediate in the plant succession — like sweetgum, say 

whereas M. ashei is a more stable component of what 
seems to be a climax association. Its extremely rapid 
growth, handsome foliage, and tolerance for drought would 
make the coastal strain a fine shade tree for use in the 
lower South. 



Magnolia pyramidata Bartr. 

This is the most common deciduous magnolia in West 
Florida. Magnolia pyramidata can be found in beech woods 
and on deciduous slopes throughout the area, both in the 
rich Knox Hill region and on the less fertile streambanks 
and bluffs along the Alabama line. Very steep north-facing 
slopes are its most typical habitat, where it can often be 
found amid mountain laurel thickets. 

Magnolia pyramidata is the coastal equivalent of M. fra- 
seri of the mountains. Whether these are separable either 
as species or as varieties is a moot question. Certainly if 
typical M. acuminata and M. acuminata var. subcordata 

* 

are to be separated, these two plants should also be. On 
occasion, the mountain plant becomes a fair-sized tree with 



72 



Rhodora t Vo1 - 77 



a thick, low-branched bole; the Florida plant is much 
smaller, with an ascending form and (usually) a single 
trunk. I do not remember ever seeing a bole bigger than 
10 cm. in diameter in West Florida. The leaves of the 
mountain plants tend to be much larger, also, though oc- 
casionally vigorous Florida specimens will belie the pub- 
lished keys. In East Texas (Jasper and Newton counties) 
putative M. pyramidata at times resembles M. fraseri in 
the size of leaves and the form and size of the trees, and is 
found, oddly enough, only on the top of a few sandy ridges ; 
no one familiar with M. pyramidata in Florida could guess 
where to seek the Texas colonies. There is a gradual transi- 
tion between M. pyramidata of Florida and M. fraseri of 
the mountains, since the Alabama and Georgia plants form 
a continuous sequence from north to south in these states. 

In Florida this plant is occasional and hardly ever pro- 
duces what might be termed colonies. The plant is by no 
means uncommon, however, and a belief that it is reflects 
not the state of nature but our limited knowledge of the 
^lexities of coastal vegetation. Like Stewartia mala- 
codendron, which is often termed rare in handbooks, M. 
miramidata seems quite frequent once its habitat is under- 



com 



stood and sought out. 



Magnolia tripetala L. 

This is apparently the first record of the occurrence of 
this magnolia in Florida. So far I have found it only along 
about half a mile of high north-facing bluff and in a con- 
tiguous ravine system on the Shoal River, approximately 
two miles west of Dorcas, Okaloosa County. This particular 
section is steep, but no more so than several other bluffs 
and ravines along the Shoal and nearby Yellow rivers. The 
bluffs are composed of Miocene marl, and the segment bear- 
ing M. tripetala does seem to be covered by an unusual 
quantity of redbud (Cercis canadensis), which in Florida 
is a good indicator of limy soil. The ravine system itself, 
where perhaps 95% of the M. tripetala plants grow, is 



1975] Magnolias — Miller 73 



a delightfully unspoiled area, with two small springfed 
streamlets running over large blocks of marl, uncut hard- 
woods such as beech and white oak and linden, an interest- 
ing herbaceous layer, and here and there the big leaves of 

ignolias. This surprising and isolated stand of this 



m 



small 



lumber 



Mag 



seems 



along Hog Creek, Randolph County, Georgia. There M 
tripetala is a vigorous competitor in the subcanopy, resem- 
bling in form and habit the specimens of M. macrophyllc 
with which it trrows: in competition with M. macrophylk 



seems 



and darkest ravines. The Florida plants, on the other hand, 



mix 



of Stewartia and small dogwood and redbud trees — the 
last of which is certainly not a remarkably tolerant species. 
I have observed no stem more than 6 or 8 cm. in diameter, 
nor is anv nlant more than 10 meters tall. The great ma- 



meters 



Mag 



ashei suggests itself, it seems likely that reduced vigor 
rather than genetic divergence accounts for the difference 
between ordinary M. tripetala and the Florida plants. For 
one thing, there are dead stems leading from a high per- 
centage of the crowns, so perhaps the leaders cannot sur- 
vive long enough to grow into a well-developed tree. The 
form of the plant is upright, unlike the horizontal tendency 
of the truly shrubby M. ashei. Most significantly, they 
show none of the ability of M. ashei to bloom vigorously at 
a small size, and that would surely be a necessary con- 
comitant to a genuine adaptation to existence as a shrub. 
In fact, no M. tripetala seedlings could be discovered in a 
rather meticulous search; only sprouts upon older root- 
stocks were discovered. The plants do show one singular 
habit, however. Approximately half the clumps are attended 
(at a distance of a meter or so) bv one or more small satel- 



74 



Rhodora [Vol. 77 



lite plants which can be traced by what appear to be hori- 
zontal runners back to the central crown. I have never 
observed this tendency to spread by vegetative means in 
other magnolias, and a check of the Hog Creek colony has 
revealed only a few sprouts which might be thought com- 
parable. At the present time I am cultivating several of 
the Florida offsets in hopes of comparing their behavior 
with a seedling M. tripetala taken from a vigorous colony 
found in coastal North Carolina near New Bern. 

Other stations close to the Shoal River colony seem to 
be (1) on the Pascagoula River in southern Mississippi 
and (2) in Butler County, Alabama (W. H. Duncan, per- 
sonal communication). Since I have, unfortunately, never 
located either station, I cannot comment on the vigor or 
habitat at other locations on the southern coastal plain. 

This deciduous magnolia seems to be the ultimate exam- 
ple of an Arcto-Tertiary relict which has persisted in an 
island of mesophytic forest on a protected north-facing 
slope amid the pinewoods of Florida. How long this colony 
has been there or whether it is the remnant of a larger 
colony would be impossible to say. For some time now it 
may even have been regenerating itself almost entirely by 
vegetative means. Its chances of surviving the chain saw 
and the log sledder and the bulldozer seem easier to esti- 
mate. It does seem unfortunate, though, that all that will 
remain of such a dogged adaptation to an alien environment 
will be a few dried sheets in a herbarium and an aberrant 
dot on a map. 8 



^Specimens of the plants discussed in this article have been de- 
posited in the herbarium of the University of Georgia. In particular, 
I would like to express my appreciation to its director, Professor 
Wilbur H. Duncan, for his unfailing generosity with his expertise 
and for his remarkable tolerance for amateur enthusiasm. 



1975] Magnolias — Miller 75 



LITERATURE CITED 

BRAUN, E. L. 1950. Deciduous forests of eastern North America. 
Hafner Publishing Co., New York. 596 pp. 

Cook, C. W., & S. Mosson. 1929. The geology of Florida. In: 
12th Ann. Rep. Florida State Geol. Survey, pp. 31-227. 

ELIAS, T. S, 1971. The genera of Fagaceae in the southeastern 
United States. Jour. Arnold Arb. 52: 150-195. 

Hardin, J. W. 1954. An analysis of variation within Magnolia 
acuminata L. Jour. Elisha Mitchell Sci. Soc. 70: 298-312. 

Harper, R. M. 1914. Geography and vegetation of northern Flori- 
da. In: 6th Ann. Rep. Florida State Geol. Survey, pp. 163-416. 

James, C. W. 1961. Endemism in Florida. Brittonia 13: 225-244. 

KURZ, H., & R. K. Goderey. 1962. Trees of northern Florida. 
University of Florida Press, Gainesville. 311 pp. 

Preston, R. J., Jr. 1961. North American trees. Iowa State Uni- 
versity Press, Ames. 395 pp. 

Sargent, C. S. 1933. Manual of the trees of North America. Ed. 2. 
Houghton Mifflin Co., Boston & New York. 934 pp. 

Thorne, R. F. 1949. Inland plants of the Gulf Coastal Plain of 
Georgia. Castanea 14: 88-97. 

Weatijerby, C. A, 1926. A new Magnolia from West Florida. 
Rhodora 28: 35-36. 



DEPARTMENT OF ENGLISH 
UNIVERSITY OF WEST FLORIDA 
PENSACOLA, FLA. 32503 






INVESTIGATIONS OF 
NEW ENGLAND MARINE ALGAE VII: 

SEASONAL OCCURRENCE 

AND REPRODUCTION OF MARINE ALGAE 

NEAR CAPE COD, MASSACHUSETTS 1 - 2 



Douglas C. Coleman and Arthur C. Mathieson 



In an earlier paper (Coleman and Mathieson, 1974) we 
described the horizontal distribution of seaweeds at seven 
sites from Scituate to Woods Hole. Massachusetts, including 



the Cape Cod Canal. In the following account we summar- 
ize the seasonal occurrence and reproduction of seaweeds at 
the same locations. Most previous collections and observa- 
tions of Cape Cod seaweeds have been restricted to the 
summer. Conover (1958) and Sears (1971) have con- 
ducted the only detailed seasonal investigations of Cape 
Cod marine algae. Conover described the productivity and 
seasonal composition of the algae in the Great Pond Estuary 
of Falmouth in relation to a variety of environmental pa- 
rameters. Sears described the subtidal benthic algae at 
several sites in southern Cape Cod. Neither Sears nor 
Conover gave any consideration to the seaweeds in the 
Cape Cod Canal, which connects Cape Cod Bay and Buz- 
zards Bay. The Canal is a transitional zone separating two 
distinctive water masses and marine floras (Coleman and 
Mathieson, 1974). The temperature discontinuity between 
the two sides of the Cape may reach 10°C. during the sum- 
mer; thus, Cape Cod is one of the major phytogeographical 
boundaries on the Atlantic Coast of North America (Far- 
low, 1870, 1882; Harvey, 1852-1858; Humm, 1969, Setchell, 
1922). 



Published with the approval of the Director of the University of 
8W Hampshire Agriculture Experiment Station as Scientific Con- 
ibution Number 665. 

2 Jackson Estuarine Laboratory Contribution No. 14. 



76 



1975] 



Cape Cod Algae — Coleman & Mathieson 



77 



Monthly collections and observations of marine algae 
Bre made at each of the seven sites (Fig. 1) during 1969. 



Specific details 



,tions, and descrip- 
summarized (Cole- 



man and Mathieson 
in the present paper. 




Figure 1. Map of Cape Cod, Massachusetts, showing the seven 
stations. 



78 



Rhodora 



[Vol. 77 



SEASONAL OCCURRENCE AND LONGEVITY 

Figures 2 and 3 summarize the monthly occurrence of 
seaweeds at each station. There was a conspicuous increase 



number 



during the spring and .summer 



except at stations 2, 3, and 4. 



time of neak numbers 



CO 

LLI 

U 

LLI 




CO 



o 




LLI 




£ 





80! 



60 



40 



20 



STATION 1 




STATION 2 




STATION 3 



40 



20 




MONTHS 

Figure 2. Monthly variation in number of species at stations 1-3. 



Black 
algae. 



red algae; hatched area 



brown algae; white 



green 



1975] 



Cape Cod Algae — Coleman & Mathieson 



79 



50 



30 



10 



STATION 4 




to 



U 




CO 



O 



oc 



CD 





STATION 5 




STATION 6 




70i 



50 



30 



10- 



STATION 7 




Figure 



o 
O. 



MONTHS 

Monthly variation in number of species at stations 4-7. 



Black = red algae; hatched area 
algae. 



brown algae; white 



green 



80 



Rhodora [Vol. 77 



varied at different stations. Seasonal changes were usually 
most obvious from January to June, except at stations 3 
and 4. A gradual decline in the number of species was 
observed from June to December at each .station. The ma- 
jority of species at each site belonged to the Rhodophyta 
and Phaeophyta ; the lowest numbers were members of the 
Chlorophyta. The red algae showed a greater increase in 
numbers during the spring and early summer than did 
either the browns or greens. Seasonal changes in numbers 
were greatest at Scituate, Wings Neck and Woods Hole. 
See Tables I-III for specific details of occurrence at stations 
1, 4 and 7 — i.e., a northern, a southern, and a Canal station. 

Of the 106 taxa collected at the seven stations, 49 were 
designated as annuals and 57 as perennials (Table IV). 
Perennials accounted for the largest number of species at 
each station, except at Wings Neck and Woods Hole. The 
ratio of annuals : perennials at each station was as follows : 
station 1 — 42 : 58, station 2 — 44 : 56, station 3 — 40 : 60, 
station 4 — 42 : 58, station 5 — 43 : 57, station 6 — 54 : 46, 
station 7 — 54 : 46. The conspicuous annuals and peren- 
nials on each side of the Cape are summarized in Tables V 
and VI. The dominance of annuals and reduced numbers of 
perennials to the south is apparent. 

Three groups of perennials can be distinguished as 
follows, depending upon their abundance north and south 
of the Cape: 1) species either restricted to or more abun- 
dant north of the Cape; 2) species either restricted to or 
more abundant south of the Cape; 3) species common on 
both sides of the Cape. The first group included Choreo- 
colax polysiphoniae, Gigartina stellata, Petrocelis midden- 
do rfii, Plumaria elegans, Fncns distichus ssp. distichus and 
Chaetomorpha atrovirens. The second group included CoZ- 
lithamnion baileyi, Sargassum filipendula, and Codium 
fragile ssp. tomentosoides. The last group included Ceram- 
ium rubrum, Chondrus crispus, Corallina officinalis, Spha- 
celaria cirrosa and Chaetomorpha linum. 

Distinct winter, spring and summer annuals were evi- 
dent at the seven sites. The first group included Bangia 



1975] 



Cape Cod Algae — Coleman & Mathieson 



81 



i l 



I 



I 

I 
I i 




ooo 




<<< 




CO CO co 




r 



I V 



T 



CO 



o 

CO 



T 



CM 



CO 



CN 



o 



T 



Q 




o 



CO 



< 



CO 






o 





< 



^ 




S3i33ds do cjggwnN 



srure 



Monthly variation of annuals at stations 1, 4 and 7 



82 



Rhodora [Vol. 77 



fmcopurpurea, Petalonia fascia and Codiohim gregarium. 
The second group included Desmarestia viridis, Desmo- 
trichum baltictim, Punctaria plantaginea, Enteromorpha 
lima, Mono stroma grevillei and Monostroma pulchrum. 
The third group included Dasya pedicellata, Champia par- 
vula, Chondria sedifolia, Grinnellia americana, Hypnea 
musciformis and Svirospora griffithsiana. Figure 4 illus- 
trates the monthly number of annual species at stations 1, 
4 and 7. The largest number of annuals occurred during 
the spring and summer months, except at station 4, and a 
decrease was evident during the winter months. 

Differential, spatial, and seasonal successions of annuals 
were recorded north and south of the Cape. Three patterns 
of seasonal succession were evident (Table VII). Some 
species occurred earlier south than north of the Cape, 
others disappeared earlier south than north, while a third 
group was found throughout the year on both sides of the 
Cape. Some examples of differential spatial succession may 



also be noted. 



fuscopu 



f 



and on the south shore of the Cape, but they were not found 
at Scituate until several weeks later. Other spring and 



summer annuals such 



dhiella 



m 



difformis appeared successive! 
stations and finally at Scituate. 



SEASONAL REPRODUCTION 



Species common to both sides of the Cape showed dif- 
ferences in reproductive periods (Tables I-III). The an- 
nuals can be divided into three groups as follows according 
to their reproductive patterns: 1) species reproducing ear- 
lier south than north of the Cape; 2) species terminating 
reproduction earlier south than north of the Cape; 3) spe- 
cies reproducing throughout the year north of the Cape, 
but with limited reproductive periods to the south. Exam- 
ples of each group are as follows: 1) Agardhiella tenera, 



1975] Cape Cod Algae — Coleman & Mathieson 88 



mar 



Chorda filum, Leathesia difformis, Ceramium strictum, 
Chordaria flag elli for mis, and Asperococcus echinatus; 2) 
Bangia, fuscopurpurea, Dumontia incrassata, Porphyra urn- 
bilicalis, Leathesia difformis, Petalonia fascia, and Mono- 
stroma grevillei; 3) Porphyra umbilicalis, Petalonia fascia, 
and Scytosiphon lomentarius. Table VIII summarizes all of 
the species found in each group. 

The perennials can also be divided into three groups 
according to their reproductive patterns: 1) species pri- 

\y reproducing during the colder months; 2) species 
primarily reproducing during the warmer months; 3) spe- 
cies reproducing throughout the year. Examples of each of 
the groups are as follows: 1) Chondrus crispus, Petrocelis 
middendorfii, Laminaria spp., and Polyides rotundas; 2) 
C all ithamni on baileyi, and Cystoclonium purpureum var. 
cirrhosum; 3) Ascophyllum nodosum, Fucus ssp., Pilayella 
littoralis, Sphacelaria cirrosa, and Ahnfeltia plicata. The 
generalized groupings of species according to reproductive 
periods were not always consistent at all stations. For 
example, Pilayella littoralis and Chondrus crispus had an 
extended reproductive period north of the Cape, but ex- 
hibited limited periods of reproduction to the south. See 
Tables I-III for specific details of reproduction at stations 
1, 4 and 7. 



DISCUSSION 



Davis (1913a, b) 



tern 



peratures near Cape Cod results in distinct annual popula- 
tions. He distinguishes two groups of annuals as follows: 
(1) winter-spring, and (2) mid-summer or early autumn. 



summer 



and aseasonal. Sears (1971) also recorded aseasonal an- 
nuals during his study of the subtidal marine algae in 
southern Cape Cod. The latter group showed no seasonal 
specificity; the plants reproduced throughout the year and 
they were represented by successive generations of young 
plants. Among others, Davis (1981a, b), Setchell (1920), 



84 Rhodora [Vol. 77 



Williams (1948) and Chapman (1964) state that species 
may survive adverse temperatures in a resting stage. 
Sears' (1971) observations substantiate the latter sugges- 
tion, for he observed germlings of several red algae over- 
wintering in the deep subtidal zone off Martha's Vineyard, 
Massachusetts. 

Cold water perennials might be expected to enter a com- 
parable state during the warmer months. Setchell (1917) 
states that the northern fucoid alga Ascophyllum nodosum 
enters a heat labor state during the summer south of the 
Cape. We have observed a similar response for A. nodosum 



become 



summer 



The ratios of annuals to perennials varied at different 
stations. North of Cape Cod perennials were dominant, 
while to the south annuals were more abundant. Inter- 



med 



tern 



regimes at the seven sites are correlated 



percentage of annuals and perennials at each site. Thus, the 
temperature range south of the Cape is about 22°C, while 
to the north it is about 17°C. ; intermediate values are evi- 
dent in the Canal (Coleman and Mathieson, 1974). The 
high summer temperatures south of the Cape allow warm 
temperature annuals to occur in abundance. In contrast the 
winter flora at the same sites is dominated by northern 
(boreal) annuals. Williams (1948, 1949) has recorded a 
similar seasonal variation of annuals at Cape Lookout, 
North Carolina. He emphasizes that areas with wide tem- 
perature fluctuations, such as Cape Lookout, support a wide 
range of annuals. 

Geographical differences of seasonal succession were evi- 
dent. For example, warm water annuals occurred earlier 



remained 



north, while many 



The 



warm 



om higher spring temperat 



than Cape Cod Bay. Likewise, some cold water annuals 
remained lonerer north of the Cane, heransp of tha lnwpr 



1975] Cape Cod Algae — Coleman & Mathieson 85 



temperatures 



ggested 



differences in seasonal successions at the stations are pri- 
marily dependent on temperature differentials. 

Variable reproductive periodicities were also evident for 



same 



exampl 



warm water annuals initiated reproduction earlier south 
than north of the Cape, while cold water annuals repro- 
duced longer north than south. The reproductive patterns 



similar natterns as 



differences 



Acknowledgements 



We would like to thank Dr. A. Hodgdon for his critical 
review of the manuscript. In addition we express our grati- 
tude to Dr. R. Fralick, Dr. R. Turner and Mr. John Hans- 
man for their assistance with diving and collection of speci- 
mens, and to the Army Corps of Engineers at Cape Cod for 
diving access to the Cape Cod Canal. 



LEGEND: TABLES Mil — a = alpha spore, b = beta spore, 
C = carpospore, G = gametang-ia, M == monospore, PS = pluriloc- 
ular sporangium, R = receptacle, S = spermatium, T = tetraspore, 
US = unilocular sporangium, Z = zoospore, X = present, — = ab- 
sent. 



86 



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1975] 



Cape Cod Algae — Coleman & Mathieson 



97 



TABLE IV. LONGEVITY OF SPECIES 

Chlorophyta 



Taxon 



Chaetomorpha atrovirens Taylor 
Chaetomorpha linum (Muller) Kutzing 



rph 



(Weber et Mohr) Kutzing 



Cladophora flexuosa (Muller) Harvey 



Cladoph 



(Griffiths ex Harvey) Kutzing 



Codiolum gr eg avium A. Braun 
Codiolum petrocelidis Kuckuck 
Codium fragile (Sur.) Hariot ssp. iomentosoides 

(van Goor) Siiva 
Enteromorpha intestinalis (L.) Link 
Enteromorpha linza (L.) J. Agardh 
Monostroina grevillci (Thuret) Whittrock 

M&nostroma pulchrum Farlow 
Rhizocloniam tortuosum Kutzing 
Spongomorpha arcta (Dillwyn) Kutzing 
Spongomorpha spinescens Kutzing 
Ulothrix flacca (Dillwyn) Thuret 

Viva lactuca L. 

Uroxpora collabens (C. Agardh) Homes et Batters 
Urospora penicilliformis (Roth) Areschoug 



Longevity 

P 
P 

P 

P? 

P? 

A 

A 



P 
A 
A 
A 
A 

P 
A 
A 
A 
A 
A 
A 



Phaeophyta 



Taxon 



Ascophyllum nodosum (L.) Le Jolis 

Asperococcus echinatus (Mertens) Greville 

Chorda filum (L.) Stackhouse 

Chorda tomcntosa Lyngbye 

Chordaria fiagelliformis (Muller) C. Agardh 

Desmarestia aeuleata (L.) Lamouroux 

Desmarestia viridis (Muller) Lamouroux 

D esmotrichum balticum Kutzing 

Desmotrichum undulatum (J. Agardh) Reinke 

Dictyosiph&n foenicvlaceus (Hudson) Greville 

Fciocarpus siUculosiis (Dillwyn) Lyngbye 

Elachista fucicola (Velley) Areschoug 

Focus distichus L. emend, ssp, distichus (C Agardh) Powell 

Fucus 



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P 
A 

A 

A 

A 

P 

A 

A 

A 

P 
A 

P 

P 



. _ mtatus (C. Agardh) Powell P 

Fucus distichus L. emend, ssp. cvancscens (C. Agardh) Powell P 



98 



Rhodora 



[Vol. 77 



Phaeophyta 
Taxon 

Fucus spiralis L. 

Fucus vesiculosus L. 

Fucus veswulosus var. sphaer ocarpus J. Agardh 

Fucus vesiculosus var. spiralis Farlow 

Giffordia granulosa 



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(Smith) Hamel 
aitzing) Batters 



Laminaria digitata (Hudson) Lamouroux 



Larninaria saccharina 



I iamouroux 



Leathesia difformis (L.) Areschoug 

Myrionema strangulans Greville 

Petalonia fascia (Muller) Kuntze 

Pilayella littoralis (L.) Kjellman 

Punctaria latifolia Greville 

Punctaria plantaginea (Roth) Greville 

Ralfsia fungiformis (Gunner) Setchell et Gardner 

Ralfsia verrucosa (Areschoug) J. Agardh 

Sargassum filipendula C. Agardh 

Scytosiphon lomentarius (Lyngbye) Link 

Sphacelaria cirrosa (Roth) C. Agardh 

Sphaerotrichia divaricata (C. AirardM Kvlin 









Longevity 

P 

P 

P 

P 

A 

A 

P 

P 

A 

P? 

A 

P 
A 
A 
P 

P 
P 

A 

P 
A 



Rhodophyceae 



Taxon 



A (j ardhiella tenera (J. Agardh) Schnntz 
Ahnfeltia plicata (Hudson) Fries 
Antithamnion americanum (Harvey) Farlow 
Bangia fuscopurpurea (Dillwyn) Lyngbye 



em 



bailey 



Callith amnion roseum (Roth) Lyngbye 
Ceramium rubrum (Hudson) C. Agardh 
Ceramium strictwm Harvey 
Champia parvula (C. Agardh) Harvey 
Cho'ndria sedi folia Harvey 
Chondrus crwpus Stackhouse 
Choreocolax polysiphoniae Reinsch. 
Clathr&morphum circumscriptum (Stromfelt) Foslie 
CoralUna officinalis L. 

Cystoclonium purpureum (Hudson) Batters var. 

drrhosum Harvpv 



Longevity 

A? 

P 

A 

A 

P 

P? 
A? 

P 
A 
A 
A 
P 
P 
P 
P 



P 



1975] Cape Cod Algae — Coleman & Mathieson 99 



Rhodophyceae (Continued) 



Taxon 
Dasya 



Longevity 



Agardh A 



Dcrmatolithon pustulatum (Lamouroux) Foslie P 

Dumontia incrassata (Muller) Lamouroux A 



(Stackhouse) 



Batters P 

Gloiosiphonia capillaris (Hudson) Carmichael ex Berkeley A 

Goniotrichum alsidii (Zanardini) Howe A 

Griflithsia tenuis C. Agardh A 

Grinnellia americana (C. Agardh) Harvey A 
Hild&nbrandia prototypus Nardo 



ormis (Wulfen") Lamou 



Melob 



urner 



Polysiphonia harveyi Bailey 
Polysiphonia lanosa (L.) Tandy 



P 
A 



Lilhophyllum corallinae (Crouan) Heydrich P 

Lithothwmniwm glaciate Kjellman P 

Lomentaria bailey ana (Harvey) Farlow A 

Lomentaria orcadensis (Harvey) Collins ex Taylor P? 



P 



Petrocelis middendorfii (Ruprecht) Kjellman P 



P 



P 



Phyllophora mcmbranifolia (Goodenough ex Woodward) 

J. Agardh 

Pliymatolithon lenormandi (Areschoug) Adey P 

Plumaria elegans (Bonnemaison) Schmitz P 

Polyides rotundas (Hudson) Greville P 
Polysiphonia denudota (Dillwyn) Greville ex Harvey 

in Hooker A 

Polysiphonia elongata (Hudson) Sprengel P 

A 

P 



Polysiphonia nigrescens (Hudson) Greville P 

Polysiphonia nor ae-n l ng line Taylor P 

Polysiphonia urceolata (Lightfoot ex Dillwyn) Greville P 

Porphyra leucosticta Thuret A 

Porphyra miniata (C. Agardh) C. Agardh A 

Porphyra umbilicalis (L.) J. Agardh A 

Rliodochorton penicillif orme (Lightfoot) Rosenvinge P 

Rhodomela confervoides (Hudson) Silva P 

P 



Rhodymenia palmata (L.) Greville 

Seirospora griffithsiana (Harvey) Dixon A 

Trailliella intricata (J. Agardh) Batters P 



100 



R hod or a 



[Vol. 77 



TABLE V. DOMINANT ANNUALS NORTH AND SOUTH 

OF THE CANAL 



ANNUALS 



North 

Bangia fuscopurpurea 
Dumontia inerassata 
Lomentaria bailey etna 
Polysiphania harveyi 
Porphyra umbilicalis 



Asperococcus echinatus 
Chorda filum 
Chorda tomentosa 
Chordaria flagelliformis 
Ectocarpus confervoides 
L eat fiesta difformis 
Petalonixt fascia 

Scytosiphon lomentaria 



Codiolum petrocelidis 

Enteromorpha intestinalis 
Mono stroma grevillei 
Mono stroma pulchrum 
Sp&ngomorpka arcta 
Spongomorpha spinescens 
Ulothrix flacca 
Viva lactuca 
Urospora collabens 
Urospora penicilliformis 



South 

Agardhiella teener a 

Antitfiamnion americanum 
Bangia fuscopurpurea 
Cadithamnion. roseum 
Ceramium strictum 
Champia parvula 

Chondria sedifolia 
Dasya pedicellata 
Dumontia inerassata 
Grinnellia americana 
Hypnea muscifortnis 
LomentaHa bailey ana 
Polysiphonia harveyi 
Porphyra umbilicalis 
Seirospora griffitfisiava 
Trailliella intricata 

Asperococcus echinatus 
Chorda filum 
Chorda tomentosa 
Chordaria flagelliformis 
r> esmotrichum undulatum 
Ectocarpus confervoides 
Leathesia difformis 

Petalonia fascia 
P une tar ia lat ifo lia 

Punctaria plantaginea 
Scytosiphon lomentaria 

Spfiaerotrichia divaricata 

Enteromorpha intestinalis 
Mono stroma grevillei 
Monostroma pulchrum 

Spongomorpha arcta 

Ulothrix flacca 
Urospora penicilliformis 



1975] 



Cape Cod Algae — Coleman & Mathieson 



101 



TABLE VI. DOMINANT PERENNIALS NORTH AND SOUTH 

OF THE CANAL 



PERENNIALS 



North 

Ahnfcltia plicata 

Ceramium rubrum 

Cliondrus crispus 
Choreocolax polysiphoniae 



South 



Callithamnion baileyi 
Ceramium rubrum 
Chondrus crispus 
Corallina officinalis 



Clathromorphum circumscriptum Cystoclonium purpureum 



Corallina officinalis 
Cystoclonium purpureum var. 

cirrhosum 
Gigartina stellata 
Hildenbrandia prototypes 

lAthophyllum corallinae 
Lithophyllum macrocarpum 

Petrocelis middendorfii 
Phyllophora membranif olia 
Phymatolithon lenormandi 
P him aria elegans 
Poly ides carpinus 
Poly siphonia lanosa 
Poly siphonia nigrescens 
Poly siphonia urceolata 
Rhodochorton penicilli forme 
Rhodomcla confervoidcs 
Rhodyynenia palmata 

Ascophyllum nodosum 
Dc&marestia aculeata 
Elachista fucicola 
Fucus distichus ssp. distichus 
Fucus distichus ssp. cdentatus 
Fucus spiralis 
Fucas vesiculosus 
Laminaria digitata 
Laminaria saccharina 
Piloiclla littoralis 
Ralfsia fungiformis 
Ralfsia verrucosa 
Sphaccla.ria cirrosa 

Chaetomorpha atrovirens 
Chaetomorpha linum 
Chaetomorpha melagonium 
Rhizoclonium tortosum 



var. cirrhosum 
Fosliella lejolisii 
Lithophyllum macrocarpum 
Phyllophora membranif olia 
Poly siphonia "nigrescens 
Poly siphonia urceolata 



Ascophyllum nodosum 
Dictyosivhon foeniculaceus 

Elachista fucicola 
Fucus vesiculosus 
var. sph aero car pits 

Pilaiella littoralis 
Sargassum filipendula 

Sphacelaria cirrosa 



Chaetomorpha linum 
Cladophora flexuosa 
Cladophora gracilis 

Codium fragile ssp. 
tomentosoides 



102 



Rhodora 



[Vol. 77 



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Cape Cod Algae — Coleman & Mathieson 



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104 



Rhodora [Vol. 77 



References 



Chapman, V. J. 1964. The Algae. Macmillan & Company, Ltd. 

London. 472 pp. 
Coleman, D. C. & A. C. Matiiieson. 1974. Investigations of New 

England Marine Algae VI. The distribution of marine algae 
near Cape Cod, Massachusetts. Rhodora 76: 537-563. 

Conover, J. T. 1958. Seasonal growth of benthic marine plants as 
related to environmental factors in an estuary. Publ. Inst. Mar. 
Sci. 5: 97-147. 

DAVIS, B. M. 1913a. General characteristics of the algal vegeta- 
tion of Buzzards Ray and Vineyard Sound in the vicinity of 
Woods Hole. Dept. Comm. and Labor, Bull. (U.S.) Bur. Fisheries 
31: 443-544. 

. 1913b. A catalogue of the marine flora of Woods Hole 



and vicinity. Dept. Comm. and Labor, Bull. (U.S.) Bur. Fisheries, 
31: 795-833. 

Farlow, W. G. 1870. XVII. List of sea-weeds or marine algae of 
the south coast of New England. U. S. Comm. of Fish and 
Fisheries. Commissioner's Report 1871-1872. Washington: Gov- 
ernment Printing Office. Pp. 281-294. 

. 1882. I. The Marine algae of New England. Ibid., 



report 1879. Pp. 1-210. 
Harvey, W. H. 1852-58. Nereis Boreali-Americana. I. Molano- 

spermae. Smithsonian Contrib. Knowl. 3: 1-150, pi 1-12. 1852; 

II. Rhodospermae. Ibid. 5: 1-258, pi 13-36. 1853; III. Chloro- 

spermae. Ibid. 10: ii. + 1-140, pi 37-50. 1858. 
HUMM, H. J. 1969. Distribution of marine algae along the Atlantic 

Coast of North America. Phycologia 7: 43-53. 
Sears, J. R. 1971. Morphology, systematics and descriptive ecology 

of the sublittoral benthic marine algae of southern Cape Cod 

and adjacent islands. Ph.D. Thesis. 295 pp. University of Mass. 
Setchell, W. A. 1917. Geographic distribution of the marine algae. 

Science 45: 197-204. 
. 1920. Stenothermy and zone-invasion. Am. Nat. 54: 



385-397. 



. 1922. Cape Cod in its relation to the marine flora of 

New England. Rhodora 24: 1-11. 
Williams, L. G. 1948. Seasonal alternation of marine floras at 

Cape Lookout, North Carolina. Am. Jour. Bot. 35: 682-695. 
. 1949. Marine algal ecology at Cape Lookout, North 



Carolina. Bull. Furman Univ. 31: 1-21. 



JACKSON ESTUARINE LABORATORY 
UNIVERSITY OF NEW HAMPSHIRE 
DURHAM, N.H. 03824 



A LIST OF THE MONOCOTYLEDONEAE OF BELIZE 

INCLUDING A HISTORICAL INTRODUCTION 

TO PLANT COLLECTING IN BELIZE 



David L. Spellman, John D. Dwyer, 

AND GERPvIT DAVIDSE 



The country of Belize, formerly British Honduras, which 
has had internal self -government since 1964, lies south of 
the Yucatan Peninsula about 800 miles south of New 
Orleans, Louisiana. The country is bounded on the north 

jxico's Territory of Quintana Roo, on the west and 



M 



by Guatemala, and on the east by the Caribbe; 
crth, the country extends from the Rio Hondo 



north to the Sarstoon River in the south, a distance of 174 



mil 



miles from 



on the sea to the western border. In area Belize is ap- 

elv the size of the State of Massachusetts. The 



oxima 



country encompasses 8,750 square miles exclusive of 116 



mile 



It is 



divided into the following political districts: Belize, Cayo, 
Corozal, Orange Walk, Stann Creek, and Toledo (Fig. 1). 
Belize is almost bisected by the Belize River. The north- 
ern half of the country is similar to the southern part of 
the Yucatan Peninsula, being a fairly level plain with a 
maximum elevation of about 120 m. above sea level. The 
southern part of the country is mountainous for the most 
part. The dominant physiographic features in the south 
are the Maya and Cockscomb Mountains. The reader is 
referred to a recent paper (Kessler et al., 1974) dealing 
with the geological history of this area. Average elevations 
in both these mountain masses lie between 850 m. and 
910 m. above sea level, with the country's highest point 



m 



ima 



though under the Koeppen classification it is classified as 
"Am" (tropical monsoon) north of the Maya Mountains 



105 



106 



Rhodora 



[Vol. 77 




Figure 1. Map of Belize showing administrative districts, and 
relationship to neighboring countries (inset). 



1975] Belize Monocotyledoneae — Spellman et al. 107 



and "Af" (tropical rainforest) in the mountainous south. 
There is a marked dry season from February to May. Pre- 
cipitation is greatest in the south and in the central moun- 
tain mass, and decreases to the north. Average annual 
precipitation ranges from 432 cm. in the Toledo District in 
the south to 132 cm. in the Corozal District in the north. 
According to Lundell (1945), in terms of area Belize has 
a floral diversity which is perhaps unequaled among the 
countries of Middle America. Standley & Record (1936) 
state that the most significant element of the flora is West 
Indian. There are a large number of plants native to Cuba, 
Jamaica, and other parts of the West Indies, which on the 
continent are known only from Belize. This provides stu- 
dents of plant geography with a unique opportunity for 
investigation. Pine stands, swamps, and marshes occur in 
the northern plain. In the south the vegetation approaches 
rainforest in comnosition. and resembles in some respects 



m 



amanian and South American elements. 

Sprague & Riley (1924) predicted a flora of about 4000 
species for the country, an estimate with which Lundell 
(1945) concurred. Standley & Record (1936) regarded 
endemism as low, citing a total of 138 species which could 
be so designated. If this be the rate of endemism, it will be 
about four percent. This is strikingly low, considering that 
Lundell estimated about fifteen percent for the entire Yuca- 
tan Peninsula. 

Sprague's statement in 1924 that the flora of Belize was 
perhaps less known than that of any other British Colony 
still holds true today. Prior to 1883, there were probably 
not more than twenty sheets of dried plants in the Kew 
Herbarium (Sprague & Riley, 1924). During the period 
1883 to 1894, this number was increased by about 200 
through the efforts of Daniel Morris, E. D. M. Hooper, 
J. Robertson, and Sir Alfred Maloney, the then Governor 

of the Colony. 

The first collection of any significant size was that of 
Morton E. Peck of the Yale University School of Forestry. 



108 



Rhodora [Vol. 77 



Professor Peck visited the country during the years 1905 
to 1907, collecting about 1000 numbers. No complete ac- 
count of the Peck collections has ever been published, but 
partial accounts have been given of the grasses by Hubbard 
(1913) and of the Dicotyledoneae by Blake (1917) and 
Sprague & Riley (1924). The Peck collections include more 
than 50 type specimens. 

Following Peck's visit to the country, virtually no bota- 
nizing occurred for two decades. A resurgence of activity 
took place in 1926, however, when a cooperative study was 
initiated by the Forestry Department of the country and 
the Yale School of Forestry, chiefly through the efforts of 
Samuel J. Record and his colleagues at Yale University. 
This led to collections by several of the University personnel 
as well as by various members of the local Forestry Depart- 
ment and employees of private timber companies. 

In 1928 and 1929 C. L. Lundell collected approximately 
700 numbers at Honey Camp in the Orange Walk District, 
under the auspices of the Tropical Research Foundation. 
In 1929, the significant collections of W. A. Schipp were 
started in Stann Creek and Toledo Districts. Schipp's bo- 
tanical activities in Belize were discussed at length by 
R. M. Lowden (1970). It appears that Schipp's sole means 
of support was obtained through the sale of herbarium and 
seed collections. His Flora of British Honduras for 1933- 
1934 is a collector's item; it is a mimeographed catalogue 
listing some 1129 flowering plants and offering for sale 
seeds and preserved plant parts. His collections from 1929 
to 1941 contain at least 3500 numbers. 

The period 1931 to 1958 is noteworthy for the quantity 
of plants collected in Belize as well as their quality. During 
the Maya Expeditions of the Carnegie Institution and the 
University of Michigan, H. H. Bartlett and C. L. Lundell 
botanized extensively in Cayo District and made smaller 
collections in Belize District (Lundell, 1940). Lundell's 
collections of 1928 through 1936 amount to about 3000 
numbers. i 



1975] Belize Monocotyledoneae — Spellman et al. 109 



During the Maya Expedition of 1931, Bartlett trained 
Percy Gentle of Belize City as a botanical collector (Bart- 
lett, 1935). Lundell later assumed the responsibility of 
directing Gentle's field activities. Gentle remained an active 
collector until shortly before his death in 1958. His col- 
lection represents the largest by a single individual, just 
under 10,000 numbers being gathered between 1931 and 

1958 (Lundell, 1960). 

The Cambridge University Expedition of 1959 and 1960 
accounts for two significant sets of collections. The first of 
these was a series of more than 700 numbers collected by 
the Expedition botanist D. R. Hunt. Hunt's collections are 
from Toledo, Stann Creek, and Cayo Districts. Most of his 
numbers seem to be from the Mountain Pine Ridge area of 
Cayo District. The second collection was made by the 
geographer of the Expedition, D. R. Stoddart. Stoddart's 
collecting activities were in conjunction with his mapping 
of certain of the cays. While his collections were not as 
extensive as those of Hunt, they are important, since they 
represent the only extensive collections from the offshore 
islands. Subsequent visits by Stoddart, with F. R. Fosberg, 
and M.-H. Sachet and D. Spellman, have yielded rather 



om 



om 



these include George Proctor of the Institute of Jamaica, 
John Dwyer of St. Louis University and the Missouri Bo- 
tanical Garden, Thomas Elias, then a graduate student at 
Southern Illinois University (Carbondale), Martin Kell- 
man of Simon Fraser University, and Edward Tyson and 
his students from Florida State University. 

The year 1970 yielded two sets of collections. The senior 
author collected about 800 numbers in January and in July 
while working with the University of Edinburgh Expedi- 
tion to British Honduras (Furley, 1972). Sidney McDaniel 
of Mississippi State University made extensive collections 

in March. 

From 1972 to the present time the bulk of plant collecting 
in Belize has been done by botanists from the Missouri 



110 



Rhodora [Vol. 77 



Botanical Garden. Thomas Croat, Curator of Phanerogams, 
collected 1794 numbers in 21 field days in the period from 
the 23rd of May to June 19, 1973. Alwyn Gentry, Assistant 
Curator at the Garden, collected about 1000 numbers during 
the same time. John Dwyer, Research Associate at the Gar- 
den, made several trips to Belize since 1972, collecting about 
2750 numbers, almost 1000 numbers being gathered at the 
same time and in the same localities visited by Croat and 
Gentry. In eleven days in the field in January, 1974, Dwyer 
returned to Belize to collect about 450 numbers in the cen- 
tral and northern sections of the country. In ten days of 
field work in January, 1974, Liesner collected 293 numbers 
in the Districts of Belize, Cayo, and Orange Walk. By 
arrangement with the Belize Government, representative 
collections from the Croat, Dwyer, Gentry, and Liesner 
lots are being selected and forwarded from the Garden for 
permanent deposit in the recently established herbarium at 
Belmopan, the new capital of Belize. 

The following are other botanists who over the past four 
years have done some collecting in Belize : 

Dr. Derek Burch, University of South Florida 
Dr. William Crankshaw, Ball State University 
Father Leonard Dieckman, S.J., St. Johns College, Belize 
City 

Dr. Richard Keating, Southern Illinois University (Ed- 

wardsville) 
Dr. Yon Linhart, University of Colorado 
Dr. Robert Long, University of South Florida 
Dr. Richard Pippen, Western Michigan University 
Dr. Paul Sorenson, Northern Illinois University 



Mario Sousa, The National University of Mexico 



Mr 



versity 



Wu 



The following list includes most of the botanists who col- 
lected in Belize during the past century or more and the 
year or years in which they collected : 



1975] Belize Monocotyledoneae — Spellman et al. Ill 



Aguilar, Mercedes 
Aitken, J. B. — 1927 
Anderson, Roger — 1970 

Bartlett, H. H. — 1931 
Brown, C.S. — 1927-1929? 

Brunt, Martin 

Burns, J. A. 

Campbell, A. J. F. — 1922 

Carrick, J. 

Castillo, A. — 1932 

Chanek, M. 

Contreras, Elias — 1968 

Crankshaw, William — 1970 

Croat, Thomas — 1973 

Day, David Fisher — 1864 

Dieckman, Rev. L. — 1970-1972 

Donald, G. H. 

Dwyer, John — 1967 ; 1972-1974 

Egler, F. E. — 1942 

Elias, Thomas — 1967 

Fralish, James — 1970 

Gentle, Percy H. — 1931-1958 

Heyder, H. M. — 1927 
Hooper, E. D. M. 

Hope, M. O. 

Hummel, C. 

Hunt, D. R. — 1959-1960 

Karling, J. S. — 1927-1928 

Keating, Richard — 1971-1972 

Kellerman, W. A. — 1906-1907 

Kellman, M. C. — 1969 

Kinloch, J. B. — 1932-1933 

Kluge, H. C — 1924? 

Lamb, A. F. A. — 1947? 

Lancaster, probably Thomas Leonard, a New Zealander 

Lazor, Robert — 1970 
Liesner, Ron — 1974 
Long, Robert — 1970 



112 Rhodora [Vol. 77 



Lundell, C. L. — 1928-1929 

McDaniel, Sidney — 1972 

Maxwell, Richard — 1967 

Meyer, William C. — 1930 

Molino, Antonio — 1954 

Maloney, Sir Alfred — 1893-1894 

Morris, Sir Daniel — 1893-1894 

Newey, Walter W. — 1970 

O'Neil, Rev. Hugh — 1936 

Peck, M. E. — 1905-1907 

Pelly, R. S. — 1933 

Pippen, R. — 1972-1973 

Proctor, G. R. — 1969 

Record, S. J. — 1926; 1930 

Robertson, Rev. J. — 1883-1884 

Rose-Innes, R. — 1970 

Sachet, M.-H. — 1972. 

Sampson, H. C. — 1928 

Schipp, William A. — 1929-1938 

Schmidt, K. P. — 1923 

Smart, H. P. — 1920; 1930? 

Sorenson, Paul — 1972 

Sousa, M. — 1973 

Spellman, D. L. — 1970-1972 

Stanton, Rev. W. A., S.J. — 1896; ca. 1906 

Stevenson, Duncan — 1927-1928 

Stevenson, Neil S. — 1927-1928 

Stocker, C. L. — 1924 
Stoddart, D. R. — 1959-1972 
Thompson, J. E. — 1927-1929; 1931 
Tyson, Edward — 1970 
Vaughn, Joseph — 1971 
Vanderveen, B. — 1973 
Winzerling, H. W. — 1926-1927 
Wunderlin, Richard — 1971 

We have been unable to determine the given names of 
the following collectors: Mrs. Armstrong; Disney; Einth; 
Espat ; and Saunders. 



1975] Belize Monocotyledoneae — Spellman et al. 113 



LIST OF THE MONOCOTYLEDONEAE 



The flora of Belize has been treated in part or in a com- 
prehensive fashion in a number of publications. Schipp's 
(1933-1934) Flora of British Honduras, Price List of Seeds 
& Herbarium Material was the first attempt to list all of 
the vascular plants from the country. This was followed 
by the checklist of Standley & Record entitled The Forests 
and Flora of British Honduras, which is considerably more 
complete. As a result of the Maya Expeditions in the 1930's, 
Lundell and other contributors published detailed taxonomic 
papers on several plant families or parts thereof. These 
appeared in two volumes entitled Botany of the Maya Area 
(Publ. Carnegie Inst. Wash., 1936, 1940). These treat- 
ments, although wider in geographical coverage than Belize, 
are a valuable source of information for the flora of Belize. 
Since 1946 the Flora of Guatemala, now nearing completion, 
has attempted to include the plants of Belize as well as those 
of Guatemala and currently represents the most authorita- 
tive source of information for the flora of Belize. 

The following list of Monocotyledoneae is presented as a 
way of updating current knowledge of this part of the 
Belize flora. The list was prepared by searching the litera- 
ture, by examining collections in herbaria, by identifying 
recently collected materials, and by checking field notes and 
unpublished lists. Any novelties encountered in studying 
very recent collections will be described and published sep- 



arately by specialists. 



made 



by botanists from the Missouri Botanical Garden are de- 
posited in the herbarium of the Garden (mo). 

We have attempted to cite individually most of the col- 
lections made in Belize since 1959, especially the extensive 
ones made by the staff of the Missouri Botanical Garden in 
the last few years. Since many of the collections made be- 
fore 1959 have been treated in the Botany of the Maya Area 
or formed a basis for the treatments in the Flora of Guate- 
mala, as well as in monographs and other taxonomic papers, 
we have not cited these specimens specifically. We have 



114 



Rhodora [Vol. 77 



examined all collections cited in this paper with the excp- 
tion of those of D. R. Hunt. 

The names of the collectors encountered in the list are 
abbreviated as follows : 
C = Thomas Croat 
D = John Dwyer 
Dieck = Leonard Dieckman 
G = Alwyn Gentry 

H = D. R. Hunt 
K = Martin Kellman 
Lies = Ron Liesner 
Lin = Yon Linhart 
Lon = Robert Long 

P = George Proctor 

Sor = Paul Sorenson 

Sp = Spellman 

V = Bruce Vanderveen 

D & L = Dwyer and Liesner 

D & P = Dwyer and Pippen 

DEM = Dwyer, Elias and Maxwell 

L & D = Liesner and Dwyer 

Sp & St = Spellman and D. R. Stoddart* 

*A11 collections by Spellman and Stoddart, cited as Sp & St, are 
from the cays. 

The authors of this paper would like to express their 
thanks to the Belize Government, especially to the Premier, 
the Hon. George Price, for his sustained interest in the 
project, as well as to Mr. E. 0. Bradley, Acting Chief 
Forest Officer, for his assistance. The extensive collecting 
done by the botanists of the Missouri Botanical Garden in 
1973-1974 has been facilitated greatly by the physical facili- 
ties provided at Ridge Lagoon Plantation, located at Mile 
IIV2 on the Northern Highway, Belize by Robert Baum- 
gartner, an American residing in Belize. The assistance 
rendered by the faculty at St. Johns College, Belize City is 
gratefully acknowledged. The senior author derived valu- 
able field experience from his association with the Univer- 



1975] Belize Monocotyledoneae — Spellman et ah 115 



sity of Edinburgh Expedition to Belize in July-August 1970 
under the direction of Dr. Peter Fuiiey. He was also fortu- 
nate in having* botanized the Cays of Belize with Dr. Ray- 
mond Fosberg and Dr. David Stoddart during June-July, 
1972. Father Leonard Dieckman, S.J., and Mrs. Lydia 
Waight of Belize City were kind enough to check the his- 
torical introduction. Special thanks are due to Dr. Thomas 
Croat for his assistance in identifying the Araceae and 
Bromeliaceae. Special thanks are due the National Geo- 
graphic Society for financial support given to Dr. Dwyer 
during the years 1973 and 1974 as well as the support re- 
ceived from the Associated Universities for International 
Education. 

In the list, those species which we judge have not been 
previously recorded from Belize in published accounts, are 
marked by an asterisk before the binomial. 

Alismataceae 

E chinodorits andrieuxii (H. & A.) Small — P 29929 

E. grandiftorus (Cham. & Schlecht.) Micheli — D 10978 

E.nymphaeaf otitis (Griseb.) Buch. 

Sagittaria lane i folia L. 
Amarvllidaceae 



Bomarea edulis (Tuss.) Herb. — P 29659 



mum 



folia (Lam.) Baker — H 



*Furcraea guatemalensis Trel. — Sor 7052 
Hippeastrum puniceum (Lam.) Urban — 
Hymenocallis cf. caribaea Herb. — 



Lin 205 



mocallis cf. caribaea Herb. — Sp 1787; Sp 1843 
H. littoralis (Jacq.) Salisb. — P 29938; Sp & St 2268; Sp 
& St 2288; Sp & St 2360; Sp & St 2449; Sp & St 2478 
Zephyranthes lindleyana Herb. 

Araceae 

Anthurium sp. — G 783 

A.aemulum Schott — D 9859; D 11560; D & L 12343; 

G8025;Spl604 
A. bakeri Hook. f. — C 24225 ; C 24453 



Q 



116 



Rhodora [Vol. 77 



A. concinnatum Schott 

A. crassinervium (Jacq.) Schott — L & D 1472 

A. denudatum Engler — C 23533 

A.gracile (Rudhe) Lindley — G 7954; V 616; V 618; 

V621 
A.scandens (Aubl.) Engler — C 23592; C 24309; D 

12690; G 7912; D 9946; P 30356 
A. scolopendrinum (Hamilt.) Kunth 
A. tetragonum Hook, ex Schott. — Sp 1948 ; V 658 

A. trinerve Miq. — G 7413 

A. verapazense Engl. — P 29825 

Dieffenbachia aurantiaca Engl. — D 11184 

D.sequina (L.) Schott 

Monstera acuminata Koch 

M. adansonii Schott — G 7434 

M. belizensis Lundell 

M. magnispatha Matuda 

M . tuberculata Lundell 
Montr ichardia, arborescens 



L & D 1644 



(L.) Schott — L & D 1448; 



Philodendron belizense Standi. 

P. brevispathum Schott — G 8423 

P. fragrantissimum (Hook.) Kunth — C 24510 

P.guttiferum Kunth — D 11306; G 8018; G 8241; P 



29700 



aceum (Jacq.) Schott — Sp 1967 



P. inequilaterum Liebm. — C 24305 ; D 11103 

P. panamense Krause — D 9891 ; G 8033 ; G 8132 

P. radiatum Schott — C 23534 ; C 24335 ; C 24512 

P.scandens C. Koch & Cello — C 23697; C 24511; D 
12718 

P. tripartitum (Jacq.) Schott — C 23670; C 24509 
Pistia stmtiotes L. — D & L 12158 

Rhodospatha acuminata C. Koch 
R. nervosa Lundell 



um 



G 



/ 



C 24332; C 24451 A 



'fo Hum 



1975] Belize Monocotyledoneae — Spellman et al. 117 



onium 



D &L 12043; G 8133; 



L & D 1468; P 29596; P 29916; Sp 1584 
Xanthosoma violaceum Schott — D & L 12078; P 29941 

Bromeliaceae 

Aechmea bracteata (Sw.) Mez — C 23922; C 23950; C 
24686; D 10116; D & L 12035; D & L 12184; P 30238 

A. brorneliae folia (Rudge) Baker 
A. dactylina Baker — G 6791 
A. mexicana Baker — D & L 12129 
A. nudicaulis L. Griseb. 

A. tillandsioides (Hart.) Baker — C 24678; P 30178; V 
607 

Ananas cornosus (L.) Merr. 

Androlepis donnell-smithii (Baker) Mez — C 23322; C 



24314 



di flora Wendl 



Catopsis aloides (Cham. & Schlecht.) Baker 
C . berteroniana (Schult.) Mez — P 29818 
C. morreniana Mez — Sp 1988 ; Sp 2002 
C. sessiliflora (R. & P.) Mez var. dioica L. B. Sm. 
Guzmania lingulata (L.) Mez — C 23768; D 9934 
Pitcairnia imbricata (Brong.) Regel 
P. hemsleyana Mez 
P. aff. integrifolia Gawl — C 23848 
P. petiolata (Koch & Bouche) Baker — P 29439 
Tillandsia balbisiana Schult. — D & L 12199; P 29529; 
P 29598 

T. bulbosa Hook. — C 24041; C 24803; D & L 12187A; 
D & L 12362 ; G 7974 ; P 29645 

T. dasyliriifolia Baker — C 23274 ; DEM 623 ; H 612 

T. dig 'data Mez 

T. excelsa Griseb. 

T. fasciculata Sw. — D & L 12167 ; Sp 1981 

T. festucoides Brong. ex Mez — C 23607 ; G 7788 ; P 30223 

T. filicifolia Cham. & Schlecht. 

T. juncea (R. & P.) Poir. — G 7961 

T. leiboldiana Schlecht. 



118 



Rhodora [Vol. 77 



monadelpha (E. Morren) Baker — G 7974 



T. polystachia L. 

T. schiedeana Steud. — C 23760 ; C 24327 ; D 9847 
T. streptophylla Scheidw. — C 23881 A; D & L 12186; 
D & L 12365 ; L & D 1600 ; P 29528 

T. sublaxa Baker 

T. utriculata L. 

T. valenzuelana A. Rich. — C 23766 ; C 23758 

T. vestita Schlecht. & Cham. 

Vriesa disticha (L.) Small 

V. paniculata ( L. ) Mez 

V. schippii L. B. Sm. 

Burmanniaceae 

Apteria aphylla (Nutt.) Barnh. ex Small 
Burmannia capitata (Walt.) Mart. — D & L 12369; P 
29684 

B. flava Mart. 

Dictyostegia campanulata Karst. 
D. orobanchioides (Hook.) Miers 
Gymnosiphon tenellus (Benth.) Urban 

Cannaceae 

Canna edulis Ker — K 451 ; P 29465 

C. indica L. — DEM 141 ; Sp 1411 

Commelinaceae 



Callisia 



H.B.K 



Campelia zanonia (L.) H.B.K. — D 9854; D 9883 
Commelina diffusa Burm.f . — K 411 
C. elegans H.B.K. 

C. erecta L. — C 23801 ; C 24100 ; G 8003 
C. longicaulis Jacq. 

Dichorisandra hexandra (Aubl.) Standi. 
Phaeosphaerion persicariae folium (DC.) C. B. Clarke 
Rhoeo spathacea (Sw.) Stearn — P 29646; Sp 1977; Sp 
1980 ; Sp 1987 ; Sp 2007 

Tradescantia belizensis Standi. 



1975] Belize Monocotyledoneae — Spellman et al. 119 



T. geniculatum Jacq. — P 30078 

T. lundellii Standi. 

Tripogawdra cumanensis (Kunth) Woodson — P 30153 

T '. g randiflora (D. Sm.) Woodson — H 399; K 592; P 

29692 ; Sp 2008 
T.serrulata (Vahl) Handlos — C 23717 ; G 7981 

Zebrina pendula Schnizl. 
Cyclanthaceae 



ensis (Mart 



palmata 



Hook 



Cyperaceae 



*Abildgaardia ovata (Burm.f.)Kral — D 11050 
Bulbostylis arenaria (Nees) Lindm. — H 189; P 29431 
B. capillaris (L.) C.B. Clarke 

B. junciformis (H.B.K.) Lindm. — H sn; H 13; H 244 
B.paradoxa (Spreng.) Lindm. — D & P 10124; H 81; 

P 29432 

B. tenuifolia (Rudge) Macbr. — C 23962 
B.vestita (Kunth) C.B. Clarke — D 11608; H 18 
Calyptrocarya glomerulata (Brong.) Urban — G 8595 
Carex polystachya, Sw. ex Vahl var. bartletti (O'Neill) 

Standi. & Steyermark — H 248 ; P 29647 
Cladlum jamaicense Crantz — C 23296; C 24995; D 

12642; DEM 629; Dieck 294; G 8527; Sp & St 2239 
Cy perns articulatus L. 

C. brevifolius (Rottb.) Endl. ex Hassk. 

C. compressus L. 

C. cyperoides (L.) Kuntze — H 283 

C. diffusus Vahl — C 24865 ; G 7997 ; G 8032 ; G 8443 

C. digitatus Roxb. — P 29926 

C. eggersii Boeck. 

C. elegans L. 

C. esculentus L. 
C. fkiVMS (Vahl) Nees 

C.haspan L. — C 23915; C 23998; C 24865; D 10756; 
H 100 ;P 29675 ;P 29716 



120 Rhodora [Vol. 77 



C. hermaphroditus (Jacq.) Standi. — C 24853 

C. humilis Kunth — P 29580 

C. imbricatus Retz. 

C. ischnos Schlecht. 

C. lanceolatus Poir. 

C. lentiginosus Millsp. & Chase 

C. ligularis L. — C 24066; C 24106; C 24533; DEM 516; 

Lin 30 ; Sp & St 2211 ; Sp 2450 
C.luzulae (L.) Retz. — C 23522; C 23800; D 12705; 

DEM 489; G 7996; G 8000; G 8431; P 29934; H 176; 

H 254 ;Sp 1591 

C. macrocephalus Liebm. — C 23370 
C. mutisii (H.B.K.) Griseb. — H 477 
C. ochraceus Vahl — C 24007 ; C 24660 ; P 29928 ; Sp 1814 
C. odoratus L. — C 24009 ; C 24579 ; G 7999 ; DEM 274 ; 
Dieck 161 ; P 29927 

C. pattens (Liebm.) Standi. & Steyermark 

C '. peruvianus (Lam.) F.N. Williams — C 24111; DEM 
515; Sp & St 2271; Sp & St 2343; Sp 2400; Sp 2469 

C. planifolius L. Rich. — Lin 1 ; Lin 2 ; Lin 31 ; Lin 71 ; 
Sp & St 2202; Sp & St 2210; Sp & St 2216; Sp & St 
2237 ; Sp & St 2257 ; Sp & St 2270 ; Sp & St 2272 ; Sp 
& St 2289 ; Sp & St 2324 ; Sp & St 2372 ; Sp & St 2385 ; 
Sp 2400 ; Sp 2408 ; Sp 2422 ; Sp 2503 ; Sp 2537 ; Sp 2560 

C.polystachyos Rottb. — C 24105; C 24865A; D 11465 

C. prolixus H.B.K. — D 10855 

C. rotundus L. — C 24085 

C. semiochraceus Boeck. 

C. Surinam ensis Rottb. — G 7996 

C. tenuis Sw. — K 419 ; K 426 

C. unioloides R. Br. — H 98 

Eleocharis caribaea (Rottb.) Blake — D & L 1255; G 

7868; DEM 607 
E. cellulosa Torr. 



E. elegans (H.B.K.) R. & S. 
259 ; H 175 

*E. elongata Chapm. — H 379 

E. filicvlmis Kunth — DEM : 



DEM 



1975] Belize Monocotyledoneae — Spellman et al. 121 



E. flavescens (Poir.) Urban 

E. inter stincta (Vahl) R. & S. 

E. maculosa (Vahl) R. & S. — E 170 ; H 415 

E. minima Kunth — H 445 

E. minutissima Britt. 

E. montana (H.B.K.) R. & S. — D & L 12328 



m 



E. nigrescens 



Vaughn et al. 276B 



fl 



Wright) C.B. Clarke — H 



Fimbristylis annua (All.) R. & S. 

F. autumnalis (L.) R. & S. 

F. castanea (Michx.) Vahl 

F. complanata (Retz.) Vahl — D 10855 

F. dichotoma (L.) Vahl — D 9525 A; H 618 

F. milacea (L.) Vahl 

F. spadicea (L.) Vahl — C 24095; D 11288; D & P 10037; 
Vaughn et al. 275 

F. spathacea Roth — Sp & St 2445 ; Sp & St 2277 ; Sp & 

St 2377; Sp 2415; Sp 2511; Sp & St 2565 
F. vahlii (Lam.) Link. 
Fuirena bulbipes Blake — C 24043; C 24125; D 11377; 



G 7877 ; G 8529 



* 



Wright — H 97 ; H 



F. incomplcta Nees 

F. simplex Vahl 

F. umbellata Rottb. — P 29579 

Hypolytrum longifolium (L. Rich.) Nees ssp. nicara- 

guense (Liebm.) Koyama — D & L 12067 
Lagenocarpus guianensis Lindl. & Nees 
Lipocarpa maculata (Michx.) Torr. 
Rhynchospora barbata (Vahl) Kunth — C 23273; C 

24082 ; D 10912A ; G 8530 ; H 342 ; H 621 ; L & D 1683 
R.cepkalotes (L.) Vahl — C 23516; C 24786; G 8411; 

H250 

R. chapmanii M. A. Curt. — DEM 442 

R. colorata (L.) H. Pfeiff. — C 23256; D 10931; G 7865; 
H393 



122 



Rhodora t Vo1 - 77 



corymbosa (L.) Britt. — C 24067 
curvula Griseb. — 



Griseb. — D & P 10050 ; H 337 

fes (Sw.) Mart. — C 23254 ; C 23341 ; C 23511 
c" 23820; C 23952; C 23963; C 24768; D 11070; D 
11482A; D 12767; D & P 10017; G 7871; G 8528 

H 169 ; H 237 ; Sp 1636 
R. divergens M. A. Curt. — C 23260 ; C 24080A ; D 10726 

D 12420 ;D 12615 
R. exaltata Kunth — D 11687 



exima 



D9050 



fascicularis (Michx.) Vahl — D 10406; D & P 10039 
qlobosa (H.B.K.) R. & S. — D 11670; D 12709; D & P 



H 



*R. globularis (Chapm.) Small — C 23966; D & P 10076; 
;H11;H309;H366 

R. hirsuta Vahl — L & D 1666 
* R . intermixta C. Wright — D 10924; D 10925; D 11601; 
D 11609; D 10473A; D & P 10038; D & P 10132A; 

G7867 
R. maris cuius Nees — H291 
R. micrantha Vahl 
R. nervosa (Vahl) Boeck. — C 23532; C 24603; D 9829; 

DEM 238; DEM 355; G 8496; H 293; K 524; Sp 1531; 

Sp 1711 
*R. oligantha Gray — H 426 

*R. plumosa Ell. — C 24036 ; DEM 651 ; D & P 10132A 
*R. podosperma C. Wright — H 196 

R.pusilla (Sw.) Griseb. 

R.radicans (Cham. & Schlecht.) H. Pfeiff. — C 23366; 

C 23399 ;D 11131 ;G 7995 
*R. rariflora (Michx.) Ell. — C 24036A 

R. robusta (Kunth) Boeck. — DEM 617 ; H 99 

R.rugosa (Vahl) Gale — D 11612; DEM 651 

R.setacea (Berg.) Boeck. — C 23277; DEM 651; L & D 
1681 

R. tenuis Link — DEM 442 
*R. torresiana Britt. & Standi. — H 230 ; P 29676 

R.watsoni (Britt.) Davidse 



1975] Belize Monocotyledoneae — Spellman et al. 123 



Scirpus chilensis Nees & Meyen 
*Schoenus nigricans L. — H 386 ; L & D 1675 
Scleria areolata Lundell 
S. bracteata Cav. — C 23294 ; C 23933 ; C 23964 ; C 24001 ; 

D 12403; D & L 12233; H 156; L & D 1424 
S. ciliata Michx. — C 23964 ; H 192 
S. eggersiana Boeck. — C 24034 A 
S.georgiana Core — C 23414; C 24080; D 10404; D 

10717; H 321; L & D 1660; L & D 1664; L & D 1676; 

L & D 1684 
S. hirtella Sw. — L & D 1679 
S. interrupta L. Rich. 
S. Mi folia Sw. — C 24508 ; C 24836 ; DEM 529 ; D & L 

12081 
S. lithosperma (L.) Sw. — C 24921; D 11013; D 12753 

S. macrophylla Presl 

S. microcarpa Nees — D 11467 ; H 494 

S. micrococca (Liebm.) Steud. — H 191 

S. mitis Berg. — DEM 470 

S. muehlenbergii Steud. — DEM 231 ; H 455 

S. pauciflora Muhl. 

S. pinctorum Britt. 

S.pterota Presl — C 23672; C 23936; C 23971; C 24034; 

C 24536; C 24943; D 11012; D 11216; D 11380; DEM 

545 ; G 8527 ; Spellman et al. 276 
S.pterota Presl var. melaleuca (Reichenb.) Uittien — H 

259 ; K 503 
S. secans (L.) Urban — C 24143; C 24475; G 8527; H 

493 ; Spellman et al. 276 
S. setacea Poir. 
S. verticillata Muhl. ex Willd. 

Dioscoreaceae 

N.B. A goodly number of very recent collections have not 
been identified (and hence not cited) due to the current 
state of confusion in the genus. 

Dioscorea bartlettii Morton — Sp 1895; Sp 1904 
D. bernoulliana Prain & Burkhill — H 267 



124 



Rhodora [Vol. 77 



D.convolvulacea Cham. & Schlecht. — Sp 1903; Sp 1918 

D. esurientium Uline 

D. macrostachya Benth. 

D. polygonoides H. & B. ex Willd. 

D. spiculiflora Hemsl. 

Eriocaulaceae 

Eriocaulon fulginosum C. Wright — H 383 

E. schiedeanum Koern. 
E. schippii Standi. 
Paeplanthus gentlei Moldenke 

P. lamarckii Kunth 

Syngonanthus gracilis (Koern.) Ruhl 

S. hondurensis Moldenke 
S. lundellianus Moldenke 

S. o'neillii Moldenke 
Tonina fluviatilis Aubl. 

Gramineae 

Acroceras zizanioides (H.B.K.) Dandy 

Andropogon bicornis L. — H 89 

A. elliotti Chapm. 

A. gerardi Vitman — Sp 1574 ; Sp 1644 

A. glomeratus (Walt.) B.S.P. — C 24044; Lin 34; Lin 

115; Lin 122; Sp & St 2210; Sp & St 2342; Sp & St 

2376 ; Sp 2389 

A. lateralis Nees 

A. leacostachyus H.B.K. — C 24006; D 11506; D 12769; 

G 7861 ; G 7866 

A. selloanus (Hack.) Hack. — H 144 ; H 430 

A. virginicus L. — Sp 1642 

Anthephora herirmphrodita (L.) Kuntze 

Aristida arizonica, Vasey 

A. capillacea Lam. — H 339 

A. divaricata H. & B. 

A. implexa Trin. — H 79 

A. toxa Cav. — L & D 1446 ; L & D 1667 

A. liebmannii Fourn. 

A. longifolia Trin. 



1975] Belize Monocotyledoneae — Spellman et al. 125 



A. orizabensis Fourn. — D 9107 ; H 419 

A. purpurasceris Poir. — G 7862 

A. recurvata H.B.K. — H 303 ; H 417 
*A. setifolia H.B.K. — D 11604 

A. tenuispica Hitchc. — C 23965; C 24081 ; D 11518 

A. ternipes Cav. 
*A. vilfifolia Henr. — D & P 10051 

Arundinella berteroniana (Schult.) Hitchc. & Chase — 

H418 
A. deppeana Nees — D & L 12229 ; H 385 

Axonopus aureus Beauv. — H 141 

A. ciliatif 'olius Swallen — H 83 

A. compressus (Sw.) Beauv. — H 441 

A. elongatus Swallen — H 490 

A. poiophyllus Chase — H 391 

A.purpusii (Mez) Chase 

A. rhizomatosus Swallen 
Bambusa vulgaris Schrad. ex Wendl. 

B. swalleniana McClure — C 23424 
Bothriochloa bladhii (Retz.) S. T. Blake — D 9921A 
Brachiaria fasciculata (Sw.) Blake — C 23453; C 23456; 

C 24045 ; C 24918 ; D 11551 ; DEM 401 ; K 421 ; Sp 1816 



B. mutica (Forsk.) Stapf — K 430 

Cenchrus brownii R. & S. — C 24980; K 435; Sp 1795 

C. echinatus L. — C 24096 

*C. incertus M.A. Curt. — Sp & St 2249 ; Sp & St 2364 
Chlorts barbata Sw. — C 24069 
C. eiliata Sw. 

C. radiata (L.) Sw. — C 23501 ; H 47 

Cynodon dactylon (L.) Pers. — Dieck 184; K 697; Sp 244 
*Dichanthium annulatum (Forsk.) Stapf — D 11181 
Digitaria cayonevisis Swallen 

D. ciliaris (Retz.) Koel. 

D. fuscescens (Presl) Henr. — D 11602 

D. horizontalis Willd. — C 23306; C 24108; Sp 1721 

D. insularis (L.) Mez ex Ekman — K 583 

D. multiflora Swallen 

Distichlis spicata (L.) Greene 



126 



Rhodora [Vol. 77 



Echinochloa colonum (L.) Link — C 23311; D 11407 
E.polystachya (H.B.K.) Hitchc. — D 10745 
* E chlnolaena gracilis Swallen — D 10424B; H 334 
Eleusine indica (L.) Gaertn. — C 23305; C 23307; C 

23311; C 23502; G 8088; K 422; D 9840 A; D 9853A; 

D 10403; Sp 1815 
Eragrostis acirti flora (H.B.K.) Nees — G 8092 
E. bahiensis Schrad. — D 9105 
E.ciliaris (L.) R. Br. — H 453; K 576; Sp & St 2971; 

Sp 2446 

E. doming ensis (Pers.) Steud. — D 11457; D & P 10034; 

D & P 10036; D 11457; D 12149; Dieck 117; H 378; 

Lin 73; Lin 116; Lin 117; Lin 119; Sp & St 2252; Sp 

& St 2378 ; Sp 2405 ; Sp 2454 
E. elliotii Wats. 

E. hirsuta (Michx.) Nees — H 468 
E.hypnoides (Lam.) B.S.P. 
E. lugens Nees 
E.maypurensis (H.B.K.) Steud. — D 10403; D & L 

12147; D&L 12212; H 194 
E. mexicana (Horn em.) Link 
E. prolifera (Sw.) Steud. 
E.tenella (L.) R. & S. 
Eriochloa punctata (L.) Desv. ex Hamilt. 
Eriochrysis cayennensis Beauv. 
Eustachys petraea ( Sw. ) Desv. — Lin 35 ; Lin 75 ; Lin 

121; Lin 163; Sp & St 2362; Sp & St 2236; Sp & St 

2250 ; Sp & St 2410 ; Sp 2437 
Gymnopogon spicatus (Spreng.) Kuntze 
Gynerium sagittatum (Aubl.) Beauv. 
Hackelochloa granularis (L.) Kuntze 
*Heniarthria cf. altissima (Poir.) Stapf. & C.E. Hubb. — 



Sp & St 2368 
Homolepis aturensis (H.B.K.) Chase — C 23634; C 
24060 ;H 101 

Hymenchne amplexicaulis (Rudge) Nees — C 23432 
Hypogynium virgatum (Desv.) Dandy — C 24192; D 
11067; D&L 12364; DEM 613; H 271; L & D 1672 



1975] Belize Monocotyledoneae — Spellman et al. 127 



Ichnanthus axillaris (Nees) Hitchc. & Chase 

/. brevivaginatus Swallen 

I. lag otis (Trin.) Swallen — H 434 

I. lanceolatus Scribn. & Smith — D 11379 

/. mexicanus Fourn. 

/. nemoralis (Schrad.) Hitchc. & Chase 

/. pollens (Sw.) Munro ex Benth. 

/. scaberrimus Swallen 

/. standleyi Hitchc. 

/. tenuis (Presl) Hitchc. & Chase 

/• villosns Swallen 

Imperata brasiliensis Trin. ■ — K 563 

/. contracta (H.B.K.) Hitchc. 

Isachne sp. — C 24019 

I.polygonoides (Lam.) Doell. 

Ischaemum latifolium (Spreng.) Kunth — C 24184; C 

24878; D 10466; D 11379; D & L 12192; L & D 1432 
Lasiacis divaricata (L.) Hitchc. var. divaricata — C 

23649; C 23707; C 3924; C 24420; D 9957; D 10971; 

D 11155; D & L 12030; D & L 12209; Sp 1713 
L. grisebachii (Nash) Hitchc. var. grisebachiz — C 



DEM 



Sp 1661 



L. oaxacensis (Steud.) Hitch, var. oaxacensis — DEM 

544 A; K 651 ;L&D 1588 
L.procerrima (Hack.) Hitchc. — C 24416; D 11194; D 

11416; D&L 12075 
L.mgelii (Griseb.) Hitchc. var. rugelii — C 24770; C 

24990; D 12757; D&L 12284; H 266; P 29950 
L. rusci folia (H.B.K.) Hitchc. var. ruscifolia — D 11005 
L.sloanei (Griseb.) Hitchc. — C 23912; K 588; Sp 1888 
L.sorghoidea (Desv.) Hitchc. & Chase var. sorghoidea 

K697 

Leersia hexandra Sw. 

LAigularis Trin. var. breviligularis (Prod.) Pyrah — C 
23496 

Leptochloa doming ensis (Jacq.) Trin 
L. fillformis (Lam.) Beauv. 



128 Rhodora [Vol. 77 



L. uninerva (Presl) Hitchc. & Chase 

L.virgata (L.) Beauv. — C 23454; K 405; Sp 1731 

Leptocoryphiurn lanatum (H.B.K.) Nees — G 7869 

Merostachys pauciflora Swallen 

Mesosetum blakei Swallen — H 346 

M.filifolium F.T. Hubb. — D 10041; D & P 10020; G 
7870 ; H 85 

Olyra latifolia L. — C 23943 ; C 24671 ; DEM 373 ; G 7847 ; 
Sp 1586 ; Sp 1659 ; Sp 1924 ; Sp 1965 

O.yucatana Chase — C 23395; C 23478; C 23604; C 
23894 ;L&D 1511 

Oplismenus burmannii (Retz.) Beauv. — K 532 

O. hirtellns (L.) Beauv. 

O. setarius (Lam.) R. & S. — P 29842 

Oryza alta Swallen 

O. latifolia Desv. 

O. saliva L. 

Panicum agrostoides Spreng. 

P. albomarginatum Nash — Sp 1631 

P. altum Hitchc. & Chase 

P. aquaticum Poir. 

P. bartlettii Swallen — C 23332 ; C 23387 ; K 551 
P. cayennense Lam. 
P. cayoense Swallen 
P. chamaelonche Trin. 

P. cijanescens Nees ex Trin. — C 24185A ; G 8051 ; H 211 ; 

H344 
P. erectifolium Nash — C 23259; C 24077; D 10716 
P. frondesceyis Mey. — D 11469 
P. furtivum Swallen — H 451 
P. fusiforme Hitchc. — H 308 ; DEM 449 
P. ghiesbreghtii Fourn. 
P. hirsutam Sw. — C 23483 ; C 23664 ; C 24285 ; K 461 

P. hirtum Lam. 

P. ichnanthoides Fourn. 

P. lanceanum Trin. 

P. lanuginosam Ell. — H 10 



1975] Belize Monocotyledoneae — Spellman et al. 129 



P.Iaxum Sw. — C 23537; C 23967; C 24025; C 24121; 

C 24185; C 24860; C 24930; D 10755; DEM 273; K 

698 ; Lin 74 ; Lin 123 ; H 442 ; H 458 
P. longUigulatum Nash 
P. lundcllii Swallen 
P. maximum Jacq. — D 12717 ; K 662 
P. milleflorum Hitchc. & Chase 
P. neuranthum Griseb. 
P. olivaceiim Hitchc. & Chase — D 9110 
P. parvifolium Lam. — H 495 
P. parviglume Hack. 
P. patulum (Scrib. & Merr.) Hitchc. 
P. pilosum Sw. — C 24880; Sp 1600; Sp 1693 
P. polycaulon Nash — H 9 

P '. poly g 'onatum Schrad. — C 24516; C 24517; G 7956 
P. pulchellum Raddi — DEM 173 
P. rudgei R. & S. — DEM 454 ; H 332 
P. selloivii Nees — H 463 
P. sphacrocarpon Ell. 
P. stagnatile Hitchc. & Chase 
P. stenodoides F.T. Hubb. 
P. tenerum Beyr. ex Trin. — H 497 
P. t rich ant hum Nees 

P. trichoides Sw. — C 24867; G 7989; G 8450; K 455 
P. tuerkheimii Hack. 
P. virgatum L. — K 687 

P. viscidellum Scribn. — C 23391; C 23398A; C 24077A; 

H253 
P. wrig htianum Scribn. — Sp 1514 

Paspalum blodgettii Chapm. — C 23939; C 24046A; H 
448 

P. botterii (Fourn.) Chase — C 23626 ; C 23668 ; C 23700 ; 
C 24769 

P. caespitosum Fliigge — C 23626; C 23668; C 23771; D 



10779 



'/ 



P. conjugatum Berg. var. conjugatum — C 23302 ; C 
24098; C 24262; C 24693: DEM 176: H 285: Sn 1666 



130 Rhodora [Vol. 77 



P. conjugatum var. pubescens Doell — Sp 1599 
P. conspersum Schrad. ex Schult. 
P. convexum H. & B. — H 195 

P. corcovadense Raddi — G 8607 ; H 440 
P. coryphaeum Trin. 
P. decumbens Sw. — C 24421 ; C 24520 
P. distichum L. — Sp 2465 ; Sp 2487 ; Sp 2509 ; Sp 2531 ; 
Sp & St 2464 

P . f asciculatum Willd. ex Fliigge — C 24130; C 24135; 
C 24183 

P.fimbriatum H.B.K. — C 23455; C 24612; DEM 242; 
S1722 

P. humb oldtianum Fliigge — H 462 

P. hngei (Fourn.) Nash — C 23828 
*P. laxum Lam. — H 442 ; H 458 ; Sp & St 2276 
*P. lineare Trin. — Sp 1513 

P.millegrana Schrad. — C 24769; D 11291; D 12469 

P. minus Fourn. — H 165 ; H 439 

P. multicaule Poir. — H 336 

P. notatum Fliigge 

P. nutans Lam. 

P. orbiculatum Poir. 

P. paniculatum L. — C 23789 ; H 154 

P. pec&w F.T. Hubb. 

P. pectinatum Nees — H 80 

P. pilosum Lam. 

P. pllcatulum Michx. — C 24033A ; C 24130 

P.pulchellum Kunth — C 23997; C 24033; D 11092; G 
7859 ; G 7864 ;H 88 ; Sp 1641 

P. reprns Berg. 

P. serpentinum Hochst. ex Steud. — C 24033 ; G 7863 

P. vaginatum Sw. — C 24047 ; C 24099 

P. virgatum L. — C 23309; C 23457; C 24107; D 9859A; 

D 11339; DEM 175; DEM 276; G 7957; Lin 36; Lin 

72 ; Lin 120 ; Lin 204A ; Sp 1732 



setosum 



complanatum (Nees) Hemsl. — DEM 
(Nees) Trin. 



1975] Belize Monocotyledoneae — Spellman et al. 131 



rus latifolius L. 
arvifolius Nash — G 8175 



Sp 



Phragmites australis (Cav.) Trin. ex Steud. — D 10383; 

H297;L&D1440 

Rhipidocladum bartlettii (McClure) McClure 

Rhynchelytrum repens (Willd.) C.E. Hubb. — L & D 1445 

Rottboellia exaltata L.f . 
Saccharum officinarum L. 
Sacciolepis myuros (Lam.) Chase 
Schizachyrium brevifolium Nees ex Kunth 
S. hirtiflorum Nees 

S. mierostachyum (Desv.) Roseng., Arr. & Izag. — D 
11206, D 12632; DEM 540 

S. semiberbe Nees — L & D 1671 

S. tenerum Nees 

Setaria scandens Schrad. ex Schult. — H 483 

S. geniculata (Lam.) Beauv. — C 24051; C 24984; Sp 

1692; Sp 1768 
S. grisebachii (Steud.) Fourn. 
S. tenacissima Schrad. ex Schult. 
S. tenax (L. Rich.) Desv. — H 421 
S. vulpiseta (Lam.) R. & S. 
Sorgastnim setosum ( Griseb. ) Hitchc. — H 20 
Sorghum halepense (L.) Pers. — DEM 650 
Spartina spartinae (Trin.) Merr. — D 11397; D & P 

10035; G 8372; Lin 168; Sp & St 2206; Sp 2291; Sp 

Sp 2562 
Sporobolus buckleyi Vasey 
S. cubensis Hitchc. — H 82 
S. indicus (L.) R. Br. — H 108 ; Sp 1730 
S. jacquemontii Kunth — C 23308; C 23875; C 24163; 

C 24992; D 9838A; D 9995A; D 11167; D 11408; D 

12431 

S.virginicus (L.) Kunth — C 24093; D 11475; Lin 4 
Lin 168; Lin 204; Sp 2182; Sp 2526; Sp & St 2234 
Sp & St 2275 ; Sp & St 2286 ; Sp & St 2382 ; Sp 2414 




Sp 2462; Sp 2508; Sp 2526; Sp 2573 
enotaphnim secundatum (Walt 



132 



Rhodora [Vol. 77 



Streptochaeta sodiroana Hack. 

Thrasya camplostachya (Hack.) Chase — H 437; H 614 

T.thrasyoides (Trin.) Chase 

Trachypogon angustifolius (H.B.K.) Nees ex Hack. — H 

86 ; H 488 
T.montufari (H.B.K.) Nees 

Triplas is purpurea (Walt.) Chapm. 

Tripsacum dactyloides L. 

T. latifolium Hitchc. — D & L 12211 ; H 315 

T. laxum Nash 

Zea mays L. 

Haemodoraceae 

Xiphidium caeruleum Aubl. 

Hy d r ochar itaceae 

Thalassia tcstudinum Konig 

Hypoxidaceae 

Hypoxis decumbens L. — H 153; Sp 1516; Sp 1629 

Iridaceae 

Cipura paludosa Aubl. — D 12801 
Marica gracilis Herb. 
Nemastylis bequaertii Standi. 

N. sylvestris (Loes) Loes (= Eustylis sylvestris Loes) 
— D & P 10064 



Sisyrinchium tinctorium H.B.K. — DEM 216; P 29857 

Juncaceae 
* J uncus marginatus Rotsk. — D 10754 

Liliaceae 

Allium cepa L. 

A. sativum L. 

Dracaena americana J. D. Sm. — D 9967; D 12602; P 

30028; Sp 1709A; Sp 1713A; Sp 1891 
Echeandia sp. — H 109 
Yucca elephantipes Reg. 

Marantaceae 

Calathea albicans Brong. 
C. altissima (P. & E.) Koern. 



1075] Belize Monocotyledoneae — Spellman et al. 133 



C. barbillana Cuf. — G 7889; G 8179; G 8479 
C. insignis Peters — C 24374 ; G 7943 ; G 8594 
C. Intea, (Aubl.) Mey. — C 24268; D 9877; 



DEM 



8034 ; P 30229 



Schum 



C . microcepkala (P. & E.) Koern. — C 24440; G 8181; 
G 8588 ; V 635 

Isehnosiphon morlaei (Eggers) Schum. — C 24400 



Mar 



DEM 



C 23874; C 24444; D 9975; 



My? 



Schum 



Stromanthe luted (Jacq.) Eichl. — D 11311 
Thdlia geniculata L. — C 23991 

Mayacaceae 

Mayaca dubletii Michx. 

M. fluvidtilis Aubl. — P 29427 

Musaceae 

Heliconia acuminata L. Rich. 

H. aurantiaca Ghiesbr. 

H. bourgdednd O.G. Peters. 

H. cham/pneiana Griggs 

H. collinsiana Griggs 

H. hirsuta L.f. var. rubriflora R.R. Sm. 



H 



1681; P 29923 



DEM 



H. mariae Hook.f. 

H. penduld Wawra 

H. psittdcorum L.f. — Sp 1963 

H. schiedcdna Kl. — D 1081 

H. spissd Griggs — P 30154 

H. cf. subul/itn R & P 



DEM 



Sp 1603 

H. vaginalis Benth. — D 9870 ; G 8144 ; G 8248 ; V 599 
H. Wdgneridna O.G. Peters. — C 24273 

Naiadaceae 

Nats guddalupensis (Spreng.) Morong 



134 Rhodora [Vol. 77 



Orchidaceae 

Bletia purpurea (Lam.) DC 
B r as savoki nodosa (L.) Lindl. 
Campy to cent intm micranthum 
C. sullivanii Fawc. & Rendle 



H 



H411 



Catasetum integerrimum Hook. — H 126 

Cattleya skinneri Batem. — H 50 

Chysis bractescens Lindl. 

Coryanthes picturata Reichenb. f. — P 30276 

C. speciosa Hook. 

Corymborchis flava Kuntze 

Cranichis sylvatica A. Rich. & Gal. — H 

Cycnoches chlorochilon Kl. 

C. warscewiczii Reichenb. f . — Sp 1952 

Diacnum bidentatum (Lindl.) Hemsl. 

Dicrium bilamellatum (Reichenb. f.) Hemsl. (= Caular 

thron bilamellatum (Reichenb. f.) Schult.) 
Dichaea tuerckheimii Schltr. 

Elleanthus linifolius Presl 

Encyclia bractescens (Lindl.) Hoehne 

E.cochleata (L.) Dressier — G 7818; Sp 1581 

Epidendrum alatum Batem. 

E. anceps Jacq. — P 30091 

E. belizense Reichenb. f . 

E. boothianum Lindl. 

E. boothii (L.) L.O. Wms. 

E. 

E. clowesii Batem. ex Lindl. 

E. cochleatum L. — G 7818 ; H 127 ; P 29905 

E. condylochilum Lehm. & Kranzl 

E. difforme Jacq. 

E. imatophyllum Lindl. — P 30275 

E. isomerum Schltr. — P 29774 

E. nocturnum Jacq. 

E. panicalatum R. & P. 

E. polyanthum Lindl. 

E. polybulbon Sw. 

E. pygmaeum Hook. 




1975] Belize Monocotyledoneae — Spellman et al. 135 



E. radiatum Lindl. 

E. rigidum Jacq. — P 29597 

E. stamfordianum Batem. 

E. stenopetalum Hook. 
E. verrucosum Sw. — P 
Erythrodes purpurea Ames 
Eulophia alata (L.) Fawc. & Rendle 




f 



batemanii 



G. baueri Lindl. 
Gongora maculata Lindl. 

G. cf . quinquenervis R. & P. — D 9939 ; G 8039 ; G 8134 
Habenaria lankesteri Ames 
H. mesodactyla Griseb. 

H. odontopetala Reichenb. f. 

H. cf. pauciflora (Lindl.) Reichenb. f. 

H. rep ens Nutt. 

H. setifera Lindl. 

I onopsis utricularioides (Sw.) Lindl. 

Isochilus crassiflorus A. Rich. & Gal. 

/. linearis (Jacq.) R. Br. var. camosiflorus (Lindl.) Correll 

Laella digbyana Benth. 

L.tibicinis (Batem. ex Lindl.) L.O. Wms. — H 327 

Ly caste cochleatam Lindl. ex Paxt. 
Macradenia brassavolae Reichenb. f. 
Masdevallia tubulifiora Ames 
Maxillaria crassifolia (Lindl.) Reichenb. f. 

M. densa Lindl. 

M. friedrichsthalii Reichenb. 

29687 
M. macleei Batem 
M. ring ens Reichenb. f. 
M. tenuifolia Lindl. — P 30188 
M. uncata Lindl. — P 29686 
Mormodcs buccinator Lindl. 
M. ring ens (Lindl.) Schlecht. 
Notylia barkeri Lindl. 
N. trisevala Lindl & Paxt. 



f._P 29774; P 29827; P 






136 Rhodora [Vol. 77 



Oncidium ascendens Lindl. 

0. carthagenense ( Jacq.) Sw. — H 132 

O. cebolleta (Jacq.) Sw. — H 137 

O. ensatum Lindl. — DEM 461 

0. luridum Lindl. 

O.pusillum (L.) Reichenb. f. — D 9938; H 410 

O. sphacelatum Lindl. — H 396? 



nithocephalus infl 
vottsiae Wats. 



Wats 



P. bnghamii Wats. — C 24156 
P. marginata Lindl. — H 136 
P . racemiflora (Sw.) Lindl. 
P. yucatanensis Ames & Schw« 
Polystachya cerea Lindl. 
P. clavata Lindl. — H 128? 



Hook 



masayens 
minor Fa^ 



Ponera striata — Lindl. 

Rhyncholaelia digbyana (Lindl.) Schltr. — H 133 

Scaphy glottis behrii (Reichenb. f.) Benth & Hook, ex 
Hemsl. 

S. cuneata Schltr. 



Batem 



may 



fragrans Lindl. 

macrantha Lindl. — DEM 460 ; H 
Spiranthes acaulis (J. E. Sm.) Cogn. 



S 



lanceolatus 



Sp 1623? 



S. orchidioides (Sw.) A. Rich. — DEM 537 
S. squamulosus Fawc. & Rendle 
S. tortilis (Sw.) L. Rich. — H 220 
Stelis ciliaris Lindl. 

Stenorrhynchos lanceolatus (Aubl.) Griseb. H 404 

Trigonidium egertonianum Batem. ex Lindl. — P 30222 
Vanilla fragrans (Salisb.) Ames 



1975] Belize Monocotyledoneae — Spellman et al. 137 



Palmae 

Acrocomia belizensis Bailey 
A. mexicana Karw. ex Mart. 

A. panamensis Bailey — C 23916 ; Sp 1832 
Asterogyne martiana Wendl. ex Hook. f. — C 24524 
Asterocaryum mexicanum Liebm. — C 23735; C 24165; 

C 24455 ;D 10906 ;G 8143 
Bactris major Jacq. (including B. balanoides (Oerst.) 
Wendl.) — C 23437 

B. trichophylla Burret— C 23972; C 24340; C 24788; D 
11101; Sp 1686 

Calyptrogyne donnell-smithii (Dammer) Burret — C 

24381 

Chamaedorea adscendens (Dammer) Burret 

C. arenbergiana Wendl. 

C. elegans Mart. 

C. ernesti-angusti Wendl. — C 23335; C 24306; D 10148; 
D & L 10907; G 7735; G 7831; G 7980; G 8129; Sp 

2005 
C. erumpens Moore 
C. geonomaeformis Wendl. 
C.lindeniana Wendl. sp. vet aff. — C 23318; C 24296; 

C 24382 ; G 7734 
C. neurochlamys Burret 
C. oblongata Mart. — G 7828 
C. schippii Burret — G 7831 A 
C.tvendlandiana Hemsl. — C 23816; C 24564; D 11107; 

D 11441; G 7854; G 7976; G 8309; G 8599 
Cocos nucifera L. 
Crysophila argentea Bartlett — C 23789A; D & L 12251; 

L & D 1621 ; Sp 1892 



C. warscewiczii (Wendl.) Bartlett, sp. vel aff. — C 23605 
Desmoncus ferox Bartlett — C 23338; C 24746; D 11352 

D. leiorhachis Burret 
D. quasillarius Bartlett 

D. schippii Burret — D 10958; G 8035; Sp 1685; Sp 1882 
Euterpe macrospadix Oerst. 



138 



Rhodora [Vol. 77 



Geonoma binerva Oerst. 
G. longipctiolata Oerst. 

G. oxycarpa Mart, (including G. mexicana Liebm.) 
C 24468; D 10921; D 11312; D & L 12096; G 7987 

Opisandra ?naya Cook. 

Orbignya cohune (Mart.) Dahlg. ex Standi. — C 23444; 

Sp 1687 
Paurotis schippii Burret 

P. wrightii (Griseb. & Wendl.) Britt. ex Britt. & Schafer 

C 23267 ; C 23953 ; D 10478 ; D 10977 ; Dwyer et al. 

365 ; G 8055 ; Sp 1617 ; Sp & St 2356 
Reinhartia gracilis (Wendl.) Burret var. gracilis 
R. gracilis var. gracilior (Burret) H.E. Moore 
R. latisecta (Wendl.) Burret 
Roystonea oleracea (Jacq.) Cook 
R.regia (H.B.K.) Cook 
Sabal may arum Bartlett 

S. morrisiana Bartlett — Sp 1893 

S. ncmatoclada Burret 

Scheelea sp. — C 23436 ; G 8596 

Schippia concolor Burret — C 24814; G 8422 

Synnechanthus fibrosvs Wendl. — C 24527; C 24556; D 

11309 ;Sp2015A 
Thrinax radiata Lodd. — Sp & St 2145 ; Sp & St 2222 ; 
Sp & St 2298; Sp & St 2357; Sp & St 2417; Sp & St 
2476; Sp & St 2557. We follow Reed (Principes 18: 



39. 1974) 



name 



'flora Sw. and T. fl 



Pontederiaceae 

Eichornia crass ipes (Mart.) Solms 
Heteranthera reniformis R. & P. — P 29942 
Pontederia lanceolata Nutt. — P 30311 
P. rotundifolia L. 

Potamogetonaceae 
Potamogeton lucens L. 
P. pcctinatus L. 
Ruppia maritima L. 



1075] Belize Monocotyledoneae — Spellman et al. 139 



Smilacaceae 



Willd. — K 488 



S 



S. lanceolate, L. 



S. lundellii Killip & Morton 

S. mexicana Griseb. ex Kimth 

S. mollis H. & B. — G 8492 ; Sp 1663 ; Sp 1878 

«S'. mvnda Killip & Morton 

S. ornata Lem. 

S. spinosa Mill. — DEM 1913 ; G 8043 

S. velutina Killip & Morton — D & L 12210 ; D & L 12275 ; 

H 436 ; P 29626 ; P 29765 

Triuridaceae 
Sciaphila picta Miers 

Typhaceae 

Typha angustifolia L. 

Xyridaceae 

Xyris ambigua Bey. ex 

X. caroliniana Walt. — 



Sp 1627 



\\ 172 



X 



DEM 



Sor 7091 ;Sor 7139 



X 



D 10407 



X. smalliana Nash — C 24015 ; D 10410; D 11610 ; D 11068 
X. subnavicularis Malme — H 199? 
X. subulata R. & P. — Dieck 101 

Zingiberaceae 

Alpinia speciosa (Wendl.) Schum. 

Costus congestus Rowlee — Sp 1966; Sp 2001 

C.pulverulentus Presl — D & L 12098; G 7731; G 7850; 

G 7910 ; G 7919 ; G 8002 ; G 8115 
C. cf. ruber Griseb. — Sp 1683; Sp 1911 ; Sp 1917 

C.spicatus (Jacq.) Sw. 

C. v illosissimus Jacq. — DEM 185 ; Sp 1684 

Hedychium coronarium Koenijr — D 11193; G 8233 

Renealmia aromatica (Aubl.) Griseb. 

R. exaltata L.f . — D 9931 ; L & D 1458 

R. mexicana Kl. 



140 Rhodora [Vol. 77 



LITERATURE CITED 

Bartlett, H. H. 1935. A method of procedure of field work in 
tropical American phytogeography based on a botanical recon- 
naissance in parts of British Honduras and the Peten forest 
of Guatemala. Misc. Paper I. Publ. Carnegie Inst. Wash. 461: 
1-25. 

Blake, S. F. 1917. Descriptions of new spermatophytes chiefly 
from the collections of Professor M. E. Peck in British Hon- 
duras. Contr. Gray Herb. 52: 59-106. 

PURLEY, P. A. (editor). 1972. University of Edinburgh Expe- 
dition to Central America 1970. General Report, vol. 1. 

Halcrow, M., & M. L. Halcrow. 1967. Orchids of Belize. 151 pp. 
The Government Printer, Belize. 

Hubbard, F. T. 1913. On the Gramineae collected by Professor 
M. E. Peck in British Honduras. 1905-1907. Proc. Am. Acad. 
Arts 49: 493-502. 

Kessler, S. E., C. F. Kienle, & J. H. Bateson. 1974. Tectonic 
significance of intrusive rocks in the Maya Mountains, British 
Honduras. Bull. Geog:r. Soc. Am. 85: 549-552. 

Lowden, R. M. 1970. William A. Schipp's botanical exnlorations in 
the Stann Creek and Toledo districts, British Honduras (1929- 
1935). Taxon 19: 831-861. 

Lttndell, C. L. 1940. The 1936 Michigan-Carnegie Botanical Expe- 
dition. Misc. Paper XIV. Publ. Carneq-ie Inst. Wash. 522: 1-57. 

— -. 1945. The vegetation and natural resources of British 



Publ 



Honduras, pp. 270-273 in F. Verdoorn (editor), Plants and Plant 
Science in Latin America. Waltham, Mass. 

. 1960. Plantae Mayanae — I: Notes on collections from 

the lowlands of Guatemala. Wrigfhtia 2: 49-63. 

Carnegie Inst. Wash. 1936. Botany of the Maya Area. Misc. 
Papers I-XIII. Publ. 461. Washington, D. C. 
. 1940. Botany of the Mnya Area. Misc. Papers XIV- 



XXI. Publ. 522. Washington, D. C. 
Schipp, W. A. 1933-1934. Flora of British Honduras, Price List 

of Seeds & Herbarium Material from William A. Schipp, Stann 

Creek, British Honduras. 
Spragite, T. A., & L. A. M. Riley. 1924. Materials for a Flora 

of British Honduras: I. Bull. Misc. Inform. 1924: 1-20. 
Standley, P. C, & S. J. Record. 1936. The Forests and Flora of 

British Honduras. Field Mus. Nat. Hist. Bot. Ser. 12: 1-432. 

DAVID L. SPELLMAN GERRIT DAVIDSE 

JOHN D. DWYER MISSOURI BOTANICAL GARDEN 

ST. LOUIS UNIVERSITY ST. LOUIS, MISSOURI 63110. 
ST. LOUIS, MISSOURI 63110. 



NOTES ON THE LEGUMINOSAE II. 
FACULTATIVE DWARFISM IN 
CROTALARIA SAGITTALIS L. 1 

Donald R. Windler- 

In his study of the North American species of Crotalaria, 
based on herbarium specimens, Senn (1939) described a 



om 



merican Southwest and Me 



name 



variety Blumeriana. During the fall of 1967, while 
studying unifoliolate species of Crotalaria in North Amer- 
ica (1970), I was able to observe a number of Mexican 
populations of this dwarf form in the field. In most loca- 
tions visited, larger plants of C. sagittalis were often found 



moi 



eemed 



ety Blumeriana. The consistency of this association \ 
to place the status of the variety in question. 

In the summer of 1968 plants reared from seeds of two 
"Blumeriana" populations (Windier & Windier 2950 — State 
of Jalisco; Windier & Windier 2981 — State of Oaxaca) 
were planted along with representatives of other species in 
a common garden plot at the North Carolina Botanical 
Gardens. A total of 47 plants, progeny of eleven seed par- 
ents from the two field populations, were grown. All plants 
grown in the garden plots were substantially larger than 
their field grown parents (see Figures 1 & 2). Plants from 
the field populations rarely grew to one decimeter in height 
and were mostly less than 0.7 dm. tall. Garden grown off- 
spring of these plants grew to heights of three to seven 
decimeters. In addition to stem length, substantial increases 
were also observed in fruit size and number of fruits per 
plant. Garden plants from the dwarf seed parents com- 



X A portion of a thesis submitted in partial fulfillment for the Ph.D. 
at the University of North Carolina at Chapel Hill. 

Contribution number 4 from the Towson State College Herbarium, 
Baltimore, Maryland 21204. Cost of publication of illustrations 
assisted by the Towson State College Faculty Research Fund. 



141 



1 42 



Rhodora 



[Vol. 77 















I 




' 






aOTAMJCAi coLtlcnom OF 









■ 



Figure 1. A. Herbarium specimen of CrotcUaria sagittaMs "var. 

Blum vri ana", B. A branch of a specimen grown from seed produced 
by the plant shown in Figure 1A. 



1975] 



Crotalaria 



Windier 



143 











A 



B 



m 






vr <- 















, 












Figure 2. A. Herbarium specimen of Crotalaria sagittalis "var. 
Blumeriana". E. A branch of a specimen grown from >(>vd produced 
by the plant shown in Figure 2A. 



144 Rhodora [Vol. 77 



pared favorably in general size with those from non-dwarf 
seed parents. 

During preparation for the garden plantings, seeds were 
scratched and soaked overnight before being placed in the 
soil. At planting time, the 47 seeds from the dwarf popula- 
tions were swollen and the radical was strongly exserted 
from the seed coat. In contrast, among the more than 1,000 
seeds from 62 other North American C rotatoria populations, 
all were swollen at planting time but none showed an ex- 



serted radicle. 



DISCUSSION 

\ information 



recognized as var. Blumeriana appear to be ecophenes of 
Mexican Crotatoria sagittalis ecotypes that are adapted to 
xeric, rocky conditions where moisture becomes available 
but is soon depleted. Under these conditions organisms 
must produce seeds which germinate quickly, and the re- 
sulting plants must quickly develop to a physiological stage 
conducive to flowering and fruit set if the population is to 
continue to exist. Plants of most other Crotatoria popula- 
tions in North America have not evolved under similar 
pressures and have not developed the quick germination 
response shown by the Mexican dwarf plants. 

The ability to germinate quickly and produce seed at a 
small plant size appear to be genetically controlled, but the 

latter characteristic is not expressed unless the plant is 
developing in a xeric situation. Consequently no formal 
recognition should be accorded to the epithet Blumeriana. 

LITERATURE CITED 

Senn, H. 1939. The North American Species of Crotalaria. Rho- 
dora 41: 317-366. 

Windler, D. R. 1967. Notes on the Leguminosae I (Neptunia and 
Cassia). The Southwestern Naturalist 12: 336-337. 

. 1970. Systematic studies in Crotalaria sagittalis and 

related species in North America (Legnminosae). Unpublished 
dissertation. University of North Carolina at Chapel Hill. 

BIOLOGY DEPARTMENT 
TOWSON STATE COLLEGE 
BALTIMORE, MD. 21204 



PECTIS HUMIFUSA NEW TO THE FLORA 

OF THE UNITED STATES 



During research preparatory to a revision of section 
Pedis of the genus Pedis L. (Compositae), a new record 
for the flora of the United States was discovered. A speci- 
men of Pedis humifusa Swartz, a taxon distributed among 
several islands of the Caribbean region, was found to have 
been collected in southern Florida. This first 
record for P. humifusa is a significant range extension, 



mai 



mi 



away in the Dominican Republic. The previously known 



from 



omimcan 



public. The following are the collection data for this new 
record: united states: Florida, Collier Co., Naples, in 



te sand along route 41, the Tamiami 

n. 24 Mar 1956. J. A. Churchill « n 



(MSC). 



Three other species of Pedis are known to occur in Flor- 
ida: P. leptocepkala (Cass.) Urban, P. linearifolia Urban 
and P. prostrata Cav. All are low herbs with yellow, five- 
rayed heads, as is P. humifusa. However, of the three pre- 



resem 



'/ 



similar in their 



mor 



prostrate habit, relatively broad leaves (usually 
2 mm. wide) and very short-peduncled or sessile heads. 
The two species differ in several respects, however, and are 
easy to distinguish. Both species tend to root at the nodes, 
but the tendency is much more developed in P. humifusa. 
The foliage of P. humifusa is blue-green and the leaves are 
mostly 4-12 mm. long and 1.5-4 mm. wide, ranging from 
oblanceolate to obovate. The leaves of P. prostrata are more 
olive-green, commonly exceed 20 mm. in length and are at 
most oblanceolate. The heads of P. humifusa are more open 
at anthesis than those of P. prostrata and are commonly 



145 



146 Rhodora [Vol. 77 



campanulate rather than cylindric or urceolate. The in- 
volucral bracts of P. humifusa are obovate, obtuse, and 
keeled only part of their length whereas those of P. pro- 
strata are usually lanceolate or oblanceolate, acute, and are 
keeled nearly to the tips. Pedis humifusa is a plant of 
sandy areas, and P. prostrata occurs in a variety of habi- 
tats and is a common roadside weed in much of Latin 
America. 



DAVID J. KEIL 
DEPARTMENT OF BOTANY 
THE OHIO STATE UNIVERSITY 
COLUMBUS, OHIO 43210 



MARITIME AND MARINE LICHENS 

FROM NAHANT 1 - - 



E. E. Webber 



The Marine Science Institute of Northeastern University, 
Nahant, Massachusetts, became operational in 1966; its first 
classes were offered during- the summer of 1967. At that 
time, Dr. Ivan Mackenzie Lamb of the Farlow Herbarium 
made an initial collection (with determinations) of the 
lichens common to the environs of the laboratory. The 
majority of lichens collected encrust the extensive outcrops 
of rock which essentially encircle this research facility. 



LICHEN DETERMINATIONS 

A. Below are listed those maritime lichens occurring on 

the metamorphic rocks above the sea: 

Acarospora fuscata (Schrad.) Arn. 
Caloplaca scopularis (Nyl.) Lett. 

( Sect. Gasparrinia ) 
Candelariella aurella (Hoffm.) Zahlbr. 
Lecanora dispersa (Pers.) Somm. 
Lecanora (Aspicilia) gibbosula Mogn. 

Lecanora rubina (Vill.) Ach. 

(Sect. Placodium) 

P armelia conspersa (Ach.) Ach. 

(Sect. XanthoparmeUa) 

Rhizocarpon grande (Florke ex Flot.) Arn. 

Rhizocarpon distlnctum Th. Fr. 

Dlmelaena or etna (Ach.) Norm. 



2 This list is published with the permission of Dr. I. M. Lamb, 
whose extreme generosity in this regard is sincerely appreciated. 

-Contribution No. 25 from the Marine Science Institute, Nahant, 
Mass. 01908. 



147 



148 



Rhodora [Vol. 77 



B. The following marine lichens appear as black, tar-like 

coatings on rock ; the first is found in the spray zone, 
while the second is abundant in the intertidal : 

Verrucaria erichsenii Zsch. 
Verrucaria mucosa Wahlenb. ex Ach. 

The above twelve lichens were collected during the course 
of one afternoon (Lamb, personal communication). Un- 



emam 



DEPT. OF BIOLOGY 

KEUKA COLLEGE 

KEUKA PARK, N.Y. 14478 



PHYCOLOGICAL STUDIES FROM THE 

MARINE SCIENCE INSTITUTE 
NAHANT, MASSACHUSETTS 1 



I. INTRODUCTION AND PRELIMINARY 
TABULATION OF SPECIES AT NAHANT 



E. E. Webber 



This is the initial paper in a series dealing with the at- 
tached marine algae at Nahant. These observations and 
studies began in the summer of 1968, and are continuing. 

The present paper summarizes the history of marine 



summai 



citations emphasizing recent research dealing with the local 

ine algal vegetation. A tabulation of the species found 



mar 



presented. No 



made at this time to discuss the plants listed since discus- 
sions of specific taxa will constitute the topics of future 
papers in this series. 



INTRODUCTION 



marine algal vegetation of New 



England began essentially in the late 1800's. The Nahant 



omin 



mtly in these studies. The efforts of 
such local investigators as Farlow, Collins, Davis, and 
others (see Taylor, 1957 for a thorough bibliography), 



om 



resulted 



1 



common 



em 



algae of the littoral and upper littoral zones of the shore- 
line. Study of New England marine algae progressed 
through the 19 20's and 1930's, culminating in the appear- 

1 Contribution No. 24 from the Marine Science Institute. I wish 
to thank Dr. A. C. Mathieson for his valuable suggestions during the 
writing of this paper. 



149 



150 



Rhodora [Vol. 77 



ance of Taylor's (1937) 



north 



North America. Twenty years later, recog- 



new 



ir both an updating of information and the 
data, Taylor published a revision (1957) 
of his volume. This major contribution is still the basic 
reference for phycologists interested in the benthic marine 
algae of our northeastern coast. 

From 1957 to the present, marine phycological studies in 
New England have progressed rapidly along several fronts. 
The utilization of SCUBA diving equipment has enabled 
the marine phycologist to study more accurately the hereto- 
fore inaccessible sublittoral zone ; in addition, this technique 
has permitted field research on a year-round basis. The 
following may be cited as recent examples of SCUBA-aided 
research in New England waters: Lamb & Zimmermann, 
1964; Hehre & Mathieson, 1970; Sears & Wilce, 1970; 
Hehre et dL, 1970; Adey, 1970; Wilce, 1970; and Mathieson, 
et al. f Nova Hedwigia, in press. 

Several important phycological discoveries have resulted 
from recent intensive field studies. For example, the ap- 
pearance of Lomentaria clavellosa in North America was 
reported from Massachusetts by Wilce & Lee, 1964; more 
recently, this species has been found in New Hampshire 
(Hehre, 1972). 

Also from New Hampshire has come the first record of 
the occurrence of H alley st is ovalls along the northeastern 
coast of North America (Mathieson & Burns, 1970). This 
raises the larger question : is there a Halicystis-Derbesia 
alternation occurring in nature among the New England 
populations of these algae, for Derbesia is common to the 
sublittoral of our area (Sears & Wilce, 1970) ? 

In addition, year-round field studies have established the 
presence of two rare and monotypic brown algal crusts 
(Petroderma ma enh forme and Porterinema fluviatile) 
known previously only from the Arctic, and now recorded 
for New England from Ipswich, Massachusetts (Wilce, 
Webber, & Sears, 1970). More specifically, this publication 



1975] Nahant Algae — Webber 151 



M 



terinema 



in North America. This latter taxon had been described 
previously only from the Baltic. For detailed discussions 
of the morphology, cytology, reproductive periodicities, and 
geographical distributions of these two "fleshy crusts", the 
reader is referred to the above paper. A comprehensive 
updating and summation of both brown and red algal 
"fleshy crusts" in New England has been presented by 
Wilce (1971). 

Complementing the renewed vigor in research on the 
algae of the sublittoral zone, recent attention also has been 
focused on estuaries and their adjacent salt marshes. In 
New England, particularly, these ecosystems represent 
promising areas of study for the marine phycologist. Pub- 
lished accounts of the benthic algal vegetation of our salt 
marshes are few; the majority of species common to such 
interesting habitats are microscopic, and many, undoubt- 
edly, have been overlooked by previous investigators. Pub- 
lications relative to New England salt marsh algae are, 
therefore, recent in their appearance. While such publica- 
tions are essentially ecologically oriented, they also deal 
with problems in systematics, life history studies, and 
polymorphisms of certain of the species encountered. The 
following papers may be mentioned as examples of recent 
studies centered about salt marsh ecosystems : Blum, I960, 
1968; Blum & Conover, 1953; Blum & Wilce, 1958; Webber, 
1967, 1968, 1971 ; Webber & Wilce, 1971, 1972; and Mathie- 
son & Fralick, 1972. The works of Drouet (1968) and 
Blum (1972), although not dealing entirely with New Eng- 
land salt marsh algae, do contain some phycological data 
applicable to this area. 

An even greater void in our understanding of the attached 
marine algae relates to the autecology of particular species. 
To date, comparatively few autecological studies have been 
undertaken. Examples of New England seaweeds in both 
field and laboratory settings include the following: Kan- 
wisher, 1957, 1966; Conover, 1958; Mathieson & Burns, 



152 



Rhodora t Vo1 - 77 



Mathieson, 1972a, 1972b; Fralick & Math 



son, 1972, 1973 ; and Jordan & Vadas, 1972. 

PRELIMINARY SPECIES TABULATION 

The species listed in the following tabulation have been 
collected and examined by me. I acknowledge the advice of 
Dr. F. Drouet concerning several of the bluegreen algal 
determinations; Dr. C. van den Hoek for assistance with 
species of Cladophora; Dr. H. W. Johansen for advice on 
the Coralline algae; and Dr. R. T. Wilce for confirmation 
of the Peyssonnelia determination. 

The systematic treatment of the Cyanophycophyta fol- 
lows that of Drouet and Daily (1956) ; Drouet (1962, 1963, 



1964, 1968); Fan (1956); and Tilden (1910). The treat- 
ments of the Chlorophycophyta, Phaeophycophyta, and Rho- 
dophycophyta all essentially follow Taylor (1957), with the 
following exceptions: van den Hoek (1963) was followed 
for Cladophora, while the recent studies of Bliding (1963, 
1968) were consulted for the Ul vales; species designations 
in Ectocarpns are after Rosenvinge & Lund (1941) and 
Cardinal (1964), Petroderma follows Waern (1952), and 
Powell's study (1957) was used for Fucus, and that of 
Wilce (1965) for Laminaria. The Chrysophycophyta (ex- 
cluding diatoms) are treated in this paper following Taylor 
(1957) for the Vaucheriales, and Parke and Dixon (1968) 
for the Ochromonadales. 

I 

Cyanophycophyta 

Anabaenatotidosa (Carm.) Lag. 
Calothrix Crustacea (Thur.) Fan 
Coccochloris stagnina Spreng. 
Entophysalis delist a (Menegh.) Dr. et D. 

Lyngbya lutea (Ag.) Gom. 

Microcoleus chthonoplastes Thur. 
M. lyngbyaceus (Kiitz.) Crouan 
M. vaginatas (Vaucher) Gom. 
Nodularia harveyana (Thwaites) Thur. 
N. spumigena Mert. 



1975] Nahant Algae — Webber 153 



Chlorophycophyta 

Eliding ia minima (Nag. ex Kiitz.) Kylin 
Brachiomonas sp. 
Bryopsis plumosa (Huds.) C. Ag. 
Capsosiphon fulvescens (C. Ag.) S. et G. 
Chaetomorpha linum (Muell.) Kiitz. 
C. melagonium (Weber & Mohr) Kiitz. 
Chlorochytrium moorei Gardner 
Cladophora albida (Huds.) Kiitz. 
Codiolum gregarium A. Br. f. intermedium 
Enter omorpha flexuosa (Wulfen ex Roth) « 
E '. intestinalis (L.) Link 
E. intestinalis (L.) Link f. clavata J. Ag. 

E. linza (L.) J. Ag. 

E. linza (L.) J. Ag. var. oblanceolata Doty 

Percursaria percursa (C. Ag.) Bory 

Pras'niocladus lubricus Kuck. 

Prasiola stipitata Suhr 

Rhizoclonium tortuosum Kiitz. 

Spong omorpha ar eta (Dillw.) Kiitz. 

S. lanosa (Roth) Kiitz. 

S. spinescens Kiitz. 

Ulothrix flacca (Dillw.) Thur. 

Ulva gigantea (Kiitz.) Bliding 



Phaeophycophyta 

Agarum cribosum (Mert.) Bory 
Alaria esculenta (L.) Grev. 
Ascophy Hum nodosum (L.) Le Jolis 
Asperococcus echinatus (Mert.) Grev. 
Chorda filum (L.) Stack. 
Chorda tomentosa Lyngbye 
Chordaria flag elli for mis (Miill.) C. Ag. 
Corynophlaea sp. 

Desmarestia aculeata (L.) Lam. 

D. aculeata (L.) Lam. var. attenuata Taylor 

D. viridis (Miill.) Lam. 



154 



Rhodora [Vol. 77 



Dictyosiphon chordaria Aresch. 



f o c niculac ens 



f 



fervoides var. confervoides (Roth) Kjellm 
/ar. siliculosus (Dillw.) Kjellm. 



/ 



ifucicola (Veil.) Aresch. 

tichus L. ssp. edentatus (de la Pyl.) Powell 



F. distichus L. ssp. evanescens (C. Ag.) Powell 
F. vesiculosus L. 



Wile 



am 



Leathesia difformis (L.) Aresch. 
Petalonia fascia (Muell.) Kuntze 

Petroderma maeuli forme (Woll.) Kuck. 

Pylaiellalitt oralis (L.) Kjellm. 

Ralfsia clavata (Harv. in Hook) Crouar 

R. fungiforms (Gunn.) S. et. G. 

R. verrucosa (Aresch.) J. Ag. 

Scy to siphon lomentaria (Lyngbye) Linl 

Sphacelaria cirrhosa (Roth) C. Ag. 

Streblonema aecidioides (Rosenv.) Fos. 



Rhodophycophyta 
Acrochaetium sp. 

Agardhiella tenera (J. Ag.) Schmitz 

Ahnf cltia plicata (Huds.) Fries 
Antithamnion floccosum (Miill.) Kleen 
Asparagopsis hamifera (Hariot) Okamura 

Bangia fuscopurpurea (Dillw.) Lyngbye 

Ceramium deslon gs champ ii Chauvin var. Hooperi (Harv.) 

Taylor 
C. rubriforme Kylin 
C. rub mm (Huds.) J. Ag. 
Chondrus crisptts Stack. 
Corallina officianalis L. 

Clathromorphum circumscriptum (Strom.) Fos. 
Cystoclonium purpureum (Huds.) Batters 



1975] Nahant Algae — Webber 155 



C. purpureum (Huds.) Batters var. cirrhosum Harv 
C . pur pur earn (Huds.) Batters f. stellatum Collins 
Damontia incrassata (Mull.) Lam. 
Euthora cristata (C. Ag.) J. Ag. 
Gigartina stellata (Stack.) Batters 



arm 



Gracilaria vemicosa (Huds.) Papenfuss 

Hildenbrandia prototypes Nardo 

Lithothamnium sp. 

Membranopteraalata (Huds.) Stack. 

Petrocelis midden dor fit (Rup.) Kjellm, 

Peyssonuelia sp. 

Phycodrys rubens (L.) Batters 

PhyUophora brodiaei (Turner) J. Ag. 

Phymatolithon laevigatum (Fos.) Fos. 

PhymatoUthon sp. 

Polyides caprinus (Gunn.) Papenfuss 

PoJysiphonia lanosa (L.) Tandy 

P.nigrescens (Huds.) Grev. 

P. novae-angliae Taylor 

P. urccolata (Lightfoot) Grev. 

Porphyranmbilicalis (L.) J. Ag. 

Ptilota serrata Ktitz. 

Rhodocorton purpureum (Lightfoot) Rosenv. 

Rhodymenia palmata (L.) Grev. 



Chrysophycophyta 

Apistonema — like plants 
Ruttnera sp. {Invar itima (Anand) Parke) 
Urococcus foslieanus Hansgr. 
Vaucheria compacta (Collins) Collins 



LITERATURE CITED 



Adey, W. H. 1970. The crustose corallines of the northwestern 
North Atlantic, including* Lithotluurinium lemoinae n. sp. Jour. 

Phycol. 6: 225-229. 
Bliding, C. 19(53. A critical survey of European taxa in Ulvales. 

I. Opera Bot. (Suppl. Bot. Not.) 8: 1-160. 



156 Rhodora [Vol. 77 



. 1968. A critical survey of European taxa in Ulvales. 

II. Ulva, Ulvaria, Monostroma, Kornmannia. Bot. Not. 121: 
535-629. 

Blum, J. L. 1960. A new Vauclteria from New England. Trans. 

Amer. Micros. Soc. 79: 298-301. 
. 1968. Salt marsh Spartinas and associated algae. Ecol. 



Monogr. 38: 199-221. 



. 1972. Vaucheriaceae. No. Am. Flora. II. 8: 1-64. 
, & J. T. Conover. 1953. New or noteworthy Vaucheriae 



from New England salt marshes. Biol. Bull. 105: 395-401. 
, & R. T. WiLCE. 1958. Description, distribution, and 



ecology of three species of Vaucheria previously unknown from 
North America. Rhodora 60: 283-288. 

Burns, R. L., & A. C. Mathieson. 1972a. Ecological studies of 
economic red algae. II. Culture studies of Chondrus cri&pus 
Stackhouse and Gigartina stellata (Stackhouse) Batters. Jour. 
Exp. Mar. Biol. Ecol. 8: 1-6. 

■ & . 1972b. Ecological studies of economic red 

algae. III. Growth and reproduction of natural and harvested 
populations of Gigartina stellata (Stackhouse) Batters in New 
Hampshire, Ibid. 9: 77-95. 

Cardinal, A. 1964. Etude sur les ectocarpacees de la Manche. Nov. 
Hedw. 15: 1-86. U figs. 

Conover, J. T. 1958. Seasonal growth of benthic marine plants as 
related to environmental factors in an estuary. Inst. Mar. Sci., 
Univ. Texas. Port Aransas 5: 97-197. 

Drouet, F. 1962. Gomont's ecophenes of the bluegreen alga Micro- 
coleus vaginatU8 (Oscillatoriaceae). Proc. Acad. Nat. Sci. 114: 
191-205. 

. 1963. Ecophenes of Schizothriz calcicola (Oscillatori- 



aceae). Ibid. 115: 261-281. 

. 1964. Ecophenes of Microcoleus chthonoplastes. Rev 



Algol. 4: 315-324. 



. 1968. Revision of the classification of the Oscillatoria- 



ceae. Monogr. 15. Acad. Nat. Sci., Philadelphia. 370 pp. 



— ■ , & W. A. Daily. 1956. Revision of the coccoid Myxo- 

phyceae. Butler Univ. Bot. Studies 12: 1-218. 

Fan, K. C 1956. Revision of the genus Calothrix Ag. Rev. Algol. 
N.S. 2: 154-178. 

Fraliok, R. A., & A. C. Mathieson. 1972. Winter fragmentation 
of Codiurn fragile (Suringar) Hariot ssp. tomentosoidcs (van 

Goor) Silva (Chlorophyceae, Siphonales) in New England. Phy- 
colo^ia. 11: 67-70. 

- & . 1973. Ecological studies of Codium fra- 



gile in New England. Mar. Biol. 19: 127-132. 



1975] Nahant Algae — Webber 157 



HEHRE, E. J. 1972. Lomentaria clavellosa (Turner) Gaillons: an 
addition to the marine algal flora of New Hampshire. Rhodora 

74: 158. 

, J. R. Conway, & R. A. Stone. 1970. Flora of the Wolf 



Islands, Part III: The marine algae. Ibid. 72: 115-118. 

, & A. C. Mathieson. 1970. Investigations of New Eng- 



land marine algae. Ill: Composition, seasonal occurrence, and 

reproductive periodicity of the marine Rhodophyceae in New 

Hampshire. Ibid. 72: 194-239. 
Jordan, A. J., & R. L. Vadas. 1972. Influence of environmental 

parameters on intraspecific variation in Fucus vexiculosus. Mar. 

Biol. 14: 248-252. 
Kanwisher, J. L. 1957. Freezing and drying in intertida] algae. 

Biol. Bull. 113: 275-285. 
. 1966. Photosynthesis and respiration in some seaweeds. 

Pp. 407-420 In: H. BARNES, (Ed.), Some Contemporary Studies 

in Marine Science. George Allen and Unwin Ltd., London. 
Lamb, I. M. & M. H. Zimmermann. 1964. Marine vegetation of 

Cape Ann, Essex County, Massachusetts. Rhodora 66: 217-254. 
Mathieson, A. C, & R. L. Burns. 1970. The discovery of Halicystis 

ovalis (Lyngbye) Areschoug in New England. Jour. Phycol. 6: 

404-405. 

. & . 1971. Ecological studies of economic red 

algae. I. Photosynthesis and respiration of Chondrus crispus 
Stackhouse and Gigartina stellata (Stackhouse) Batters. Jour. 
Mar. Biol. Ecol. 7: 197-206. 

, & R. A. FRALICK. 1972. Investigations of New England 

marine algae. V. The algal vegetation of the Hampton-Seabrook 
Estuary and the open coast near Hampton, N. H. Rhodora 74: 
406-435. 
, E, J. Hehre, & N. B. Reynolds. Investigations of New 



England Marine Algae. I. A floristic and descriptive ecological 
study of the marine algae at Jaffney Point, New Hampshire. 
Nov. Hedw. (in press). 
, N. B. Reynolds, & E. J. Hehre. Investigations of New 



England marine algae. II. The species composition, distribution 
and zonation of seaweeds in the Great Bay Estuary System and 
the adjacent open coast of New Hampshire. Ibid, (in press). 

Parke, M., & P. S. Dixon. 1008. Check-list of British marine algae 
— second revision. Jour. Mai'. Bio. Assoc. U. K. 48: 783-832. 

Powell, H. T. 1957. Studies in the genus Fucus L. I. Fucus 
distichus L. emend. Powell. Ibid. 36: 407-432. 

ROSENVINGE, L. K., & S. LUND. 1941. The marine algae of Denmark. 

Contributions to thair natural history. II. Phaeophyceae. Part 
I. T). Kg]. Danske Vidensk., Biol. Skrifter., Bd. 1. Kobenhavn. 



158 Rhodora [Vol. 77 



Sears, J. R., & R. T. Wilce. 1970. Reproduction and systematics 
of the marine alga Derbesia (Chlorophyceae) in New England. 
Jour. Phycol. 6: 381-392. 

Taylor, \V. R. 1937. Marine Algae of the Northeast Coast of 
North America, ix + 427 pp. Univ. Mich. Press. Ann Arbor. 



1957. Marine Algae of the Northeast Coast of North 
America. (Revised ed.). viii + 509 pp. Univ Mich. Press. Ann 
Arbor. 

Tilden, J. 1910. Minnesota Algae. I. Myxophyceae of North 

America. Minneapolis. 
Van den Hoek, C. 1963. Revision of the European species of 

Cladophora. E. J. Brill. Leiden. 248 pp. + 55 pb. 
Waern, M. 1952. Rocky-shore algae in the Oregrund Archipelago. 

Acta Phytogeogr. Suecica 30: 1-298. Uppsala. 
Webber, E. E. 1967. Rluegreen algae from a Massachusetts salt 

marsh. Bull. Torrey Bot. Club 94: 99-106. 
■ . 1968. New England salt marsh Vaucheriae. Rhodora 



70: 274-277. 



. 1971. Observations on Microcolcux lynghyaceus (Kiitz.) 

C rowan from marine habitats in New England. Ibid. 73: 238- 
243. 

, & R. T. Wilce. 1971. Benthic salt marsh algae at 

Ipswich, Massachusetts. Ibid. 73: 262-291. 



& . 1972. The ecology of benthic salt marsh 

algae at Ipswich, Massachusetts. I. Zonation and distribution 
of algal species. Ibid. 74: 475-488. 

Wilce, R. T. 1965. Studies in the genus Laminaria. Pp. 247-256 
In: Bot. Gothoburgensia. III. Proc. Fifth Mar. Biol. Symposium. 



Goteborg. 



1970. Cladophora pyffinaea Reinke in North America. 



Jour. Phycol. 6: 260-263. 



. 1971. Some remarks on the benthic chrysophytes and 

tho fleshy red and brown crusts. Pp. 17-25 In: Symp. Cold 
Water Inshore Marine Biology — Some Regional Aspects. Mar. 
Sci. Inst., Nahant, Mass. 

, & R. W. Lee. 1964. Lvmentaria clavellosa in North 

America. Bot. Mar. 6: 251-258. 

, E. E. Webber, & J. R. Sears. 1970. Petroderma and 

Porterinema in the New World. Mar. Biol. 5: 119-135. 



DEPT. OF BIOLOGY 
KEUKA COLLEGE 
KEUKA PARK 
NEW YORK 14478 



PHRAGMITES COMMUNIS IN SOUTH CAROLINA 



Phragmites communis Trinius, the tall reed, is a coarse 
rhizomatous perennial reed forming extensive colonies in 
open marshy habitats in almost every state of the eastern 
United States. It occurs sporadically in the southeastern 
states but has not been previously recorded in South Caro- 
lina or Georgia (Radford et al., 1968). It does occur 
sporadically in North Carolina, favoring a few of the 
northeastern and southeastern counties. 

A large colony of Phragmites was discovered by the 
author in Georgetown County, South Carolina, covering 
an area of several hundred square meters. The colony is 
located approximately 100 meters west of the Waccamaw 
River (at high tide) and 125 meters north of Route 17 
South. Vouchers of immature specimens were collected and 
are on deposit at the herbarium of the University of South 
Carolina, Columbia, South Carolina. 



LITERATURE CITED 



Radford, A. E., H. E. Ahles, & C. R. Reel. 1968. Manual of the 
vascular flora of the Carol inas. 1183 pp. Univ. North Carolina 
Press, Chapel Hill. 



RICHARD STALTER 
DEPARTMENT OF BIOLOGY 
ST. JOHN'S UNIVERSITY 
JAMAICA, NEW YORK 11432 



159 



BUTOMACEAE: A NEW FAMILY RECORD 

FOR NORTH DAKOTA 



Butomus umbellatus L., flowering rush, was collected on 
July 25, 1972, four miles south of Valley City, North Da- 
kota (SEVi Sec. 9, R58W, T139N). Scattered colonies in 
flower were found growing in shallow water along the 
shoreline and on silted gravel bars of the Sheyenne River 
for a distance of about four miles (God f read 4640, 4882, 
and 4926, NDA). These stations were revisited in 1973 and 
1974 and the colonies were increasing in size. 

Butomus umbellatus is a native of Eurasia but has be- 
come established along the St. Lawrence River Valley. 
Herbarium records indicate that the species existed in Que- 
bec in 1916. Rousseau (1966) indicates that B. umbellatus 
and another Eurasian introduction, Ly thrum salicaria L., 
purple loosestrife, are common along the shoreline of the 
St. Lawrence River, appearing to be as much in equilibrium 
as if in their native habitat. It is interesting to note that 
L. salicaria is also well established along the Sheyenne 
River near Valley City. 

Since its introduction into North America, Butomus 
umbellatus has become established at widely scattered lo- 
calities. In 1956 it was found in Idaho. It was collected in 
Wisconsin for the first time in 1957, and it is known from 
Flathead County, Montana (Hahn, 1973). In 1972 it was 
reported from Rice County, Minnesota, while Van Bruggen 
(1972) reports B. umbellatus in South Dakota from Faulk 
County, 150 miles south of the North Dakota plants along 
the Sheyenne River. The establishment of B. umbellatus in 
North Dakota is apparently rather recent. Mr. and Mrs. 
Russell Pederson, who own adjacent farmland, first noticed 
the plants five or six years ago. 

Literature Cited 

Hahn, R. E. 1973. Flora of Montana Conifers and Monocots. De- 
partment of Botany, Montana State University, Bozeman, Mon- 



tana. 143 pp. 



160 



1975] Butomaceae — Godfread & Barker 161 



Rousseau, J. 1966. Movement of plants under the influence of man. 
Pp. 81-89. hi: K. L. Taylor and R. A. Ludwig [ed.], The Evo- 
lution of Canada's Flora. University of Toronto Press, Toronto. 

Van Bruggen, T. 1972. A Manual for the Identification of the 
Vascular Plants of South Dakota. University of South Dakota 
Educational Media, Vermillion, South Dakota. 46.") pp. 



CAROLYN GODFREAD 

WILLIAM T. BARKER 

DEPARTMENT OF BOTANY 

NORTH DAKOTA STATE UNIVERSITY 

FARGO, N.D. 58102 



GERANIUM SIBIRICUM L. (GERANIACEAE) 

IN WISCONSIN 



This species, adventive from Eurasia, has heretofore been 
reported to extend from New York and Pennsylvania to 
Illinois (Fernald, Gray's Manual, ed. 8, 1950.). I report 
here its discovery in Wisconsin, from records in the her- 
baria at wis and osh. The curators of the herbaria at mil 
and UWM have kindly informed me that they have no rec- 
ords of the plant from Wisconsin. The four collections 
listed below from two counties warrant the inclusion of 
the plant as an element in the Wisconsin flora. 

Wisconsin : DANE CO. : large colony along town road close 
to village of Dane, H. C. Greene, 17 Aug. 1954 (wis) ; on 
disturbed sandy peat in grazed area of south Lake Wabesa 
marsh, Don Samuelson, 5 Sept. 1972 (wis) ; in shady, damp 
soil along Elm Drive, U. of Wisconsin-Madison Campus, 
Thomas 002, 3 Sept. 1972 (wis). COLUMBIA CO.: along a 
sandy roadside on Chryslaw Road, to the NW. of the junc- 
tion of State Route 113 and county trunk J, Sect. 19, R7E, 
TION, Zaudtke 053, 9 Sept. 1972 (osh). 



NEIL A. HARR1MAN 

DEPARTMENT OF BIOLOGY 

U. OF WISCONSIN — OSHKOSH 

54901 



162 



EDITOR'S NOTE TO CONTRIBUTORS 



manuscript 



m 



a substantial backlog- of papers awaiting review and a lag 
time approaching twenty-one months between time of re- 
ceipt and time of publication. The Editorial Committee 
consider this an unreasonable length of time and wish to 
return to a schedule allowing publication within approxi- 
mately one year of receipt. 

Rising costs on every hand, among other factors, prevent 
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indirectly to the plants of the Western Hemisphere. How- 
ever, floristic notes, range extensions, and descriptions of 
formae are discouraged unless they are of considerable 
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funds. 



mem 



M 



s 



163 



164 



Rhodora [Vol. 77 



Address manuscripts, proofs, and related correspondence 



to: 



Dr. A. Linn Bogle 
Editor-in-Chief, Rhodora 
Botany & Plant Pathology Dept 
University of New Hampshire 
Durham, N. H. 03824 



Volume 77. No. 809, including panes 1-1<>4, teas issued May SO, 1975. 



Cover illustration 

Rhododendron canadense (L.) Torr. 

from 
Curtis' Botanical Magazine t.474 



RHODORA March, 1975 Vol- 77, No. 809 



CONTENTS 



1 



41 



On the Epibiotic and Pelagic Chlorophyceae, Phaeophyceae, and 
Rhodophyceae of the Western Sargasso Sea 

William J. Woelkerling 

Saxifrages on Mount Washington 

Frederic L. Steele 

Rudbeckia auriculata (Perdue) Krai, a Species Distinct from 
R. fulgida Ait. 

Robert Krai 44 

Apparent Ecotypic Differences in the Water Relations of Some 
Northern Bog Ericaceae 
Peter J. Marchamd 53 

The Deciduous Magnolias of West Florida 

Ronald F. Miller 



r>4 



ions of New England Marine Algae VII: Seasonal 



Massachusetts 



Reproduction of Marine Algae Near Cape Cod 



Douglas C. Coleman and Arthur C. Mathieson 7fl 

A List of the Monocotyledoneae of Belize Including a Historical 
Introduction to Plant Collecting in Belize 

David L. Spellman, John D. Dwyer, and Gerrit Davidse 105 

Notes on the Leguminosae II: Facultative Dwarfism in Crotalaria 
sagittalis L. 

Donald R. Windier 141 

Pectis humifusa New to the Flora of the United States 

David J. Keil 145 

Maritime and Marine Lichens from Nahant 

E. E. Webber 

Phycological Studies from the Marine Science Institute, Nahant, 
Massachusetts I: Introduction and Preliminary Tabulation of 
Species at Nahant 

E. E. Webber 149 

Phragmites communis in South Carolina 

Richard Stalter 

Butomaceae: A New Family Record for North Dakota 



147 



159 



Carolyn Godfread and William T. Barker lfiO 

Geranium sibiricum L. (Geraniaceae) in Wisconsin 

Neil A. Harriman 



162 



Editor's Note to Contributors iflg 




JOURNAL OF THE 



NEW ENGLAND BOTANICAL CLUB 




ol. 77 



June, 1975 



No. 810 



Wc\t 5fofa ^ngkith botanical (ttlub, £0nc. 



Botanical Museum, Oxford Street, Cambridge, Mass. 02138 



Conducted and published for the Club, by 
ALFRED LINN BOGLE, Editor-in-Chief 

ROLLA MILTON TRYON 

STEPHEN ALAN SPONGBERG 

GERALD JOSEPH GASTONY 
RICHARD EDWIN WEAVER 



Associate Editors 



RHODORA. — A quarterly journal of botany, devoted primarily to 
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be 



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Forms may be closed five weeks in advance of publication. Ex- 
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Address manuscripts and proofs to: 

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University of New Hampshire, 
Durham, New Hampshire 03824 

Subscriptions and orders for back issues (making all remittances 
payable to RHODORA) should be sent to RHODORA, Botanical 
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JOURNAL OF THE 



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Vo1 - 77 June, 1975 No. 810 



DE PLANTIS TOXICARIIS E MUNDO NOVO 

TROPICALE COMMENTATIONES XII 



Notes 



Richard Evans Schultes 



Recent field work has served to corroborate the suspicion 
long- held by botanists that the Piperaceae represent a fam- 
ily of plants of very basic importance in the ethnopharma- 
cology of primitive societies of the New World tropics. 
The following notes are offered as a contribution to our 
growing knowledge of the use of piperaceous species for 
their variety of biodynamic activity. 

C. De Candolle, in DC. Prodr. 



emai 



16, pt. 1 (1869) 437. 
COLOMBIA: Comisaria del Putumayo, Mocoa and 
vicinity. December 8, 1942. R. E. Schultes et C. E. Smith 
3028. 

Called in the Ingano language cung amanda-amb e , this 
herb "pounded and mixed with tobacco and urine" is used 
"to poultice bites of the cungamanda ant". 

Peperomia glabella (Sw.) A. Dietrich var. melanostigma 

Dahlstedt, in Kgl. Sv. Vet. Akad. Handl. 33, 
pt. 2 (1900) 122. 

COLOMBIA: Comisaria del Putumayo, Mocoa and 
vicinity. Alt. 820 m. "For mal de ojo". December 6, 1942. 



R. E. Schultes et C. E. Smith 2059. 



165 



166 



Rhodora t VoL 77 



This herb is a supposed remedy for conjunctivitis. In 



Mocoa 



gwtnan. 



Peperomia serpens (Sw.) Loudon, Hort. Brit. (1830) 13. 

COLOMBIA: Comisaria del Putumayo, Rio Sucum- 
bios, Santa Rosa. "Remedy for bite of conga ant. Kofan 

: u-nu-se' -he-pa". April 7-8, 1942. R. E. Schultes 3589. 

i aromatic leaves and stems of Peperomia serpens are 



name 



em 



ant. In Kofan, the term se -he-pa refers to a medicinal or 
poisonous plant. 

Piper AHenii Trelease, in Ann. Mo. Bot. Gard. 25 (1938) 826. 

PANAMA: Provincia del Darien, trail between Pin- 
ogana and Yavisa. Altitude about 15 m. March 17, 1937. 
P. A. Allen 270. 

Allen reports that the "roots are used by Indians to 
deaden pain" and the leaves "as a snake bite remedy". 

Piper auritum Humboldt, Bonpland et Kunth, Nov. Gen. et 

Sp. 1 (1815) 54. 
EL SALVADOR: Vicinity of San Salvador. Alti- 
tude 650-850 m. February 2-7, 1922. P. C. Standley 20550. 
According to Standley, the juice of the crushed leaves of 
this highly aromatic species is employed to remove ticks. 

tVip shmh i« lorallv railed Santa Maria. 



Piper Bartlingianum C. De Candolle, in L»e ^anaone rroai . 

16, pt. 1 (1869) 257. 
DUTCH GUIANA: Fetikruk. On sandy hills. Au- 
gust 10, 1939. Geykes sine num. 

The collector reports Piper Bartlingianum as an ingredi- 
ent of "oerali poison" amongst the Wayana Indians. 

Geykes sine num. appears to represent the same species 
as A. C. Smith 2826 and 2827 from British Guiana. It is 



elements of Waiwai 



dian arrow poison. 



1975 1 Piperaceae — Schultes 167 



dactylostigmum Yuncker, in Inst. Bot. Sac 

No. 3, (1966) 35, fig. 29. 

BRAZIL: Estado do Amazonas, Manaos 



W 



ity, Reserva Ducke. "Climber, a< 
Leaves and stem rapidly numb ti 
April 13, 1972. R, E. Schultes et 

The rapid, strong and long lasting numbness of the 
tongue produced when the leaves and stems of this species 
are chewed is well known to the inhabitants of the forested 
areas around Manaos. So far as could be ascertained, how- 
ever, this property does not constitute the basis of any 
medicinal or other folk use of Piper daetylostigmum. 
Numbing of the tongue and mucous membranes of the 
mouth, which, though not common, is known for other 
species of the genus, sometimes leads to local medicinal 
use. The leaves and twigs of Piper corcovadensis (Miq.) 
DC. and P. Jaborandi Veil., for example, are chewed in the 
region of Rio de Janeiro to relieve toothache because of 
their strong numbing effect (Mors, W. B. and C. T. Rizini : 
Useful plants of Brazil (1966) 89). 



Piper erythroxyloides R. E. Schultes et Garcia-Barriga sp. 



nov. 



Frutex glaber, usque ad V/ 2 ped. altus, erectus. Ram 



rami 



gracilibus elongatisque, subgranulosis, glabris, 4.5-6 cm. 
longis. Folia membranacea. ellintica. a.nW 



acum 



aequilater subrotundata, 14-18 cm. longa, 6-8.5 cm. lata, 
omnino pinnatim venosa, venis primariis glabris, utrinque 
8-10, supra glabra, subtus minute punctulosa et irregu- 
lariter albo-squamulosa; petiolo subcarnosulo, usque ad 1 
cm. longo, longitudinaliter striato, scobiculato. Pendunculus 
subcarnosulus, usque ad 1 cm. longus, glabrus vel minutis- 
sime albido-papilloso-pilosus, bracteis crassis, suborbicu- 
laribus, cupulatisque, plus minusve 1 mm. longis. Drupa 
carnosa, extus papillosa, conica, 2 mm. in diametro, 1 mm. 

longa, in stylum crassum, 0.75 mm. lontrum desinens. 



168 



Rhodora f Vo1 - 77 



COLOMBIA: Departamento de Santander del Norte, 
La Motilonia, Rio Catatumbo y Rio Brandy. Altitude 80 m. 
"0.5 m. alto, erecto. Amentos verde-oscuros. Al masticar 
el tallo se le siente im piquante fuerte y luego se duerme la 
lengua y los labios. Los indios bari (motilones) lo mastican 
frecuentemente como la coca. Por lo tanto es narcotico". 
May 24-26, 1965. Hernando Garcia-Barriga et Gustavo 



Lozano-G 18414. 



Colomb 



Ames 



Piper erythroxyloides appears to be most closely allied to 
P. dariense C. DC. of northern Colombia and Panama. It 
differs in various important respects. The leaves of Piper 
erythroxyloides are basally rounded and apically short 
acuminate, with a blunt tip (not cuneate and long acumi- 
nate with a sharp tip) ; the internodes are not so slender 
and are somewhat longer; the inflorescence is usually 
longer, and thicker; the flowers are more congested, not 
loosely arranged ; the style is thicker ; and the stigmas are 
relativelv longer. Perhaps the most noticeable difference 
lies in the shape of the drupe : in Piper erythroxyloides, the 
fruit is conic and rough-papillose, whereas in P. dariense 
it is globose-tetragonous and either smooth or obscurely 
papillose. 



According to 



10, the stems of 
Motilone (Bari) 



moutJ 
name 



Indians as a kind of narcotic which has local effects in the 

that resemble those of coca: whence the specific 
erythroxyloides. Upon mastication, the sterns induce 
a strong burning sensation which is followed by numbness 
of the tongue and lips. 

In addition to this use, the plant is chewed in the belief 
that it prevents dental caries. 

Local Indian names of Piper erythroxyloides are achi- 
kaira and chanquira. 



note that in February, 1971, Mr 



M. Ajami of Ha 
Museum several 



1975] Piperaceae — Schultes 169 

incomplete material can indicate, to represent the same 
species: Piper erythroxyloides. According to Mr. Ajami, 
the Bari Indians of the central Venezuelan region west of 
Lake Maracaibo, who chew the internodes as an aphrodisiac, 
experience a strong analgesic effect in the mouth and con- 
spicuous dilation of the pupils. Inasmuch as the tribe is 
apparently the same group as the Colombian "Motilones" 
and the two localities are not distant, the information com- 
municated by Mr. Ajami assumes greater significance. It 
is hoped that complete botanical specimens from the Vene- 
zuelan locality may be forthcoming. 

Piper hispidum Swartz, Prodr. Veg. Ind. Occ. (1788) 15. 

ECUADOR : Parroquia Concepion, Playa Rica. Alti- 
tude 91 m. "Forest near stream; undergrowth. Perennial 
herb 4 m. high; aments erect. Common". December 6, 
1936. Y. Mexia S407. 

The collector reports that the leaves of this plant are 
"crushed in water to kill head lice". It is locally known as 

pipilongo. This collection is the type of Piper pediculicidum 
Trelease. 



Mus 



Bot. 13, no. 357 (1936) 176. 



PERU: Departamento de Loreto, alto Rio Purus, 
Zapote. "Culina Indian name : tetsi. Substitute of tobacco 
snuff. The leaves are dried and reduced into powders. Tree 
about 10 m. high. Lowland forest". October 15, 1968. Lau- 
rent Rivier 21. 

This interesting report of the preparation of a snuff from 
the leaves of a species of Piper — a snuff used as a "substi- 
tute" for tobacco — provides a significant addition to our 
growing understanding of plants employed in South Amer- 
ica in the preparation of powders for inhalation. The 



num 



much 



pected, and, in this case, the indication of psychoactive 
constituents — probably essential oils — adds to the ethno- 
pharmacological interest of the report. 



170 



Rhodora [Vol. 77 



Piper sp. 

BRAZIL: Estado do Amazonas, Rio Livramento, 
Humayta, near Livramento. October 12-November 6, 1934. 
B. A. Krukoff 6972. 

The stem of this vine-like species of Piper, according to 
the collector's notes, contains "a substance producing local 
anaesthesis". The plant is employed locally by the Indians 
"to cure toothache" and is called cipo de dor dente ("tooth- 
ache vine"). 

Pothomorphe umbellata (L.) Miquel, Comm. Phyt. (1840) 

36. 
COLOMBIA: Comisaria del Putumayo, Rio Guam- 
ues, San Antonio. Secondary growth near path. "Kofan 
Indian name: a-nama-he se' -he-pa. Curare, used alone or 



mixed ; for m 
stem and roo 



uita). Bark of lower 
Herb 5-6 feet". Sep- 



tember 5, 1966, H. V. Pinkleij 421. Same locality. Febru- 
ary 18, 1966, Pinkley 119. 

The preparation of arrow poisons amongst the Kofan 
Indians of the border region of Colombia and Ecuador is 
ethnobotanically extremely complex. Although piperaceous 
plants are known to be employed as ingredients of arrow- 
poisons over a wide area in tropical America (Hegnauer, 
R. : C hemotaxonomie der Pflanzen 5:321. 1969), Pinkley's 
report of the utilization of Pothomorphe umbellata "alone 
or mixed" assumes special significance. Most, if not all, of 
the piperaceous species employed in these preparations are 
not known to have curare constituents, and their use as 



may 



If a 



species be utilized "alone" — that is, as the only ingredient 
in a curare — it must have a biodynamically active con- 
stituent. 



CHROMOSOME COUNTS OF COMPOSITAE 
FROM THE UNITED STATES, MEXICO, 

AND GUATEMALA 1 



David J. Keil and Tod F. 'Stuessy 



Chromosome numbers can be extremely useful in system- 
atic studies, particularly for helping to reveal evolutionary 
relationships. For the past fifteen years numerous chromo- 

plants have been published, especially 
in the Compositae, and these counts have been compiled 
in several major sources (Darlington & Wylie, 1955; Cave, 



some reports from 



Moor 



Howeve 



not only that many species never have been counted, but 
also that many taxa are known only from a single plant in 
one population. In view of the common occurrence of 
euploid and aneuploid races in plants as illustrated by 
several detailed investigations (e.g., Lewis, 1962, 1970; 
Stuessy, 1971a), it is desirable to have several to 



m 



from 



m 



Kovanda, 1972; Strother, 1972). The present paper helps 

ies in the Compositae by: (1) re- 



em 



chromosome 



and varieties; and (2) reporting additional populational 



mosome 



MATERIALS AND PROCEDURES 

The meiotic chromosomal material for this study was 
collected during the past several years by the senior and 
junior authors on various field excursions. Immature capit- 
ula were killed and fixed in modified Carnoy's fluid (4 
chloroform: 3 absolute alcohol: 1 glacial acetic acid) and 
refrigerated in the laboratory until later prepared by con- 



1 Publication No. 843 from the Department of Botany, The Ohio 



State University, Columbus. 



171 



172 



Rhodora [Vol. 77 



acetocarmine squash techniques. Voucher speci- 



mens 



herbarium of The Ohio State University (OS) ; vouchers 



urn 



of Texas at Austin (TEX) . 



RESULTS 

The chromosome counts obtained in the present study 
are listed in Table 1. First counts are reported for two 
genera, 16 additional species, and one variety; 112 addi- 
tional counts are for taxa counted previously, seven of 
which are new numbers. The first counts for genera are 
from Epaltes Cass. (?t=10) and Tricarpha Longpre 
(n = S), and first counts for species are in Bidens L., 
Calea L., Guardiola Cerv. ex H. & B., Machaeranthera 
Nees, Mekimpodium L., Otopappus Benth., Sclerocarpus 
Jacq., Scnecio L., Sigesbeckia L., Simsia Pers., Spilanthes 
Jacq.. Tridax L.. and Zaluzania Pers. 



DISCUSSION 

Because many of the counts presented here corroborate 
previous chromosomal reports, the discussions are restricted 
either to first counts or to new reports for genera, species, 
or varieties. The order of commentary will follow the se- 
quence of tribes in the classification of Hoffmann (1890-94), 
which is the same as that used in Table 1. References for 
statements regarding the range of chromosomal variation 
within genera will not be given; documentation for these 
counts comes from the several major sources cited in the 
introduction to this paper. 

eupatorieae. Counts for three herbaceous species of 
Stevia Cav. represent new reports. Stevia elatior H.B.K. 
is cited here as n = 12n & 12i (Fig. 1), whereas the two 
previously recorded numbers have been n = 34i (Powell & 
Turner, 1963) and n = 33i (Grashoff, Bierner, & North- 
ington, 1972). Our count for Stevia origanoides H.B.K., 



1975] Chromosome Counts — Keil & Stuessy 173 



reported here as n = 11 (Fig. 2), is the first for this taxon 
at what appears to be the diploid level ; the previous counts 
were n -- 34i and n = ca. 43 ± li (Grashoff et al., 1972). 
Stevia plummerae A. Gray var. durang crisis Robins, has 
been reported before only once by Grashoff et al. (1972) as 
n = ca. 17, but our count is n — 44i (Fig. 3). As pointed 
out by Grashoff et al. (1972), it is common to find varying 
meiotic chromosomal associations and numbers in species 
that have apomictic races, as are present in these three 
taxa. It is not surprising, therefore, that our reported 
counts add to this chromosomal diversity. 

astereae. Several previous counts have been reported 
for Erigeron karwinskianus DC: 2n = 32 (Carano, 1924; 
Battaglia, 1950) ; 2n = 36 (Fagerlind, 1947; Larsen, 1953, 
1954; Kliphuis & Wieffering, 1972); n = 9 and n = 27i 
(Turner, Ellison, & King, 1961) ; and n — ca. 27 (Turner, 
Powell, & King, 1962). Considering the variation in chro- 
mosome number that has been documented previously in this 
species, as well as our new report of n = 5n & 17i (Fig. 4), 
it is likely that E. karwinskianus is apomictic through at 
least part of its range from Mexico to northern South 
America (Solbrig, 1962). It is interesting that our count 
comes from a population very near the locality cited by 
Turner et al (1961) for their counts of n = 9 and n = 27i. 

Machaeranthera coulteri (A. Gray) Turner & Home (as 
Psilactis coulteri A. Gray) was reported as n = 5 by Sol- 
brig, Anderson, Kyhos, Raven, and Riidenberg (1964). 
However, based on the recent revision of sect. Psilactis of 
Machaeranthera by Turner and Home (1964), the geo- 
graphic location of the voucher for the count seems more 
appropriate for M. arida Turner & Home than for M. 
coulteri. The latter species, as recently interpreted, is 
known only from the vicinity of Guaymas, Sonora, 
where our voucher was collected. Our first count of n = 5 
(Fig. 5) for Machaeranthera coulteri is particularly inter- 
esting because in the previously mentioned revision of sect. 
Psilactis of the genus (Turner & Home, 1964; cf. their 



174 



Rhodora 



[Vol. 77 








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Figs. 1-22. Camera lucida drawings of meiotic chromosomes of 
species of Compositae. Diplotene, Fig. 18; diakinesis, Figs. 5, 7-10, 
12, 15, 19-21; metaphase I, Figs. 1, 3, 4, 6, 11, 14, 16, 17; metaphase 
II, Figs. 2 (one half of cell shown), 13. All figures same scale. 
Bivalents black, univalents white. KC = Keil & Canne, KM 
& McGill, K — Keil. Fig. 1, Stevia elatior, K 9396, n 
Fig. 2, Stevia ori</a'noides, KC 8884, n 
var. durangensis, KC 8927-1, n 



Keil 



12n & 12i; 

11; Fig. 3, Stevia plummerae 

aricinskianus, 



KC 9178 n = 5n & 17i; Pig. 5, Machaeranthcra coulteri, KC 8637, 

m *-w -w— « « mm r t * J 



n 



5; Fig. 6, E patten mexicana, KC 9211, w 



10; Fig. 7, fiir/ras 



1975] 



Chromosome Counts 



Keil & Stuessy 



175 



% • 





I 






• 



%% M 







V + 







/ 







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/> 























20 



'+' 






5m 



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11 



riparia var. refraeta, KC 8710, ro = 12; Fig. 8, Cuardiola platyphyUa, 
KM 8558, n = 12; Fig. 9, Mclampodium append iculatum , KC 870GA, 
n = 10; Fig. 10, Otopappus imbricatus, KC 9112, w = 16; Fig. 11, 
Parthenium incanum, KM 7765A, 'n 
carpus spatulatv 
KC 8902, n = 



3 



18n & 18i; Fig. 12, Sclero- 
= 11; Fig. 13, Sigesbeckia jornllensix, 

= 17; 



0; Fig". 14, Simsia eurylcpis, KC 9231, n 



17; Fig-. 16, Spilanthes phane- 



Fig. 15, Simsia grayi, KC 9081, w 

rac^z',5, /fC 9035, n — 41; Fig. 17, Tricarpha durangensis, KC 8860 A, 
n = 8; Fig*. 18, Tridax tenui folia var. microccphala, KC 8808, n = 9; 
Fig. 19, Zaluzania grayana, KM 8379 A, n = 17; Fig-. 20, Schkuhria 

10; Fig. 21, Senecto 



pinnata var. guatemalensis , K 9402A, n 

runcinatus, KC 9192, n = 22; Fig. 22, Pinaropappiis roseus, KC 9177, 



n 



20jj & It. 



176 



Rhodora [™. 77 



M 



in the x 



M. arida 



M. crispa (Brandg.) Turner & Home, both known chromo- 

5. More recently M. arizonica Jackson & 



soma 



M. par v if. 



to this group and both specie^ have been co » nt€ ^f ^~ jj 
(Jackson & Johnson, 1967). * ' " L -^-~ 



chromosomal 



LVX M co^en substantiates its phyletic association with 
these other species. All other taxa in sect. Psilactis are 
known chromosomally as either n = 4 or n - 9 (Turner 
&Horne, 1964). 

INULEAE. The first report for Epaltes (E. 



mexicana 



soma 



,. — 10 (Fig 6), is in keeping with its present subtribal 
disposition in the Plucheinae. Of the related genera of the 
same subtribe (Hoffmann, 1890-94) that are known chromo- 
o— ^ (Blumea DC, Pluchea Cass., Ptengeron (DC.) 
Benth., Pterocaulon Ell., Sphaeranthus L., and Tessarra 
Ruiz & Pav.) , all are based on x = 10 except Blumea which 
appears multibasic with x = 9, 10, and 11. On morphologi- 
cal and geographical grounds, in our opinion, Epaltes mexi- 
cana Less, is quite similar to some species of Pluchea, the 
former differing mainly in its smaller heads and flowers 
and in its epappose achenes. As emphasized by Bentham 
(1873) and Godfrey (1952), the generic boundaries in the 
Plucheinae are not well defined and perhaps should be re- 
evaluated. 

HELIANTHEAE. Bidens Hparia is reported for the first 



lme 



most 



;ed numbers of » = 12, 24, and 36 (clearly based on 

x = 12). _ 

The first count for Calea zacatechichi Schlecht, n — ca. 

19 is in keeping with previous reports for other species of 
the genus (n = 9, 16, 18, 19, 24, 32). According to the 
most recent revision of the Mexican and Central American 
taxa (Robinson & Greenman, 1896), C. zacatechichi is most 
closely related to C. nelsonii Robins. & Greenm. which has 
been counted as n = ca. 18 (Turner et al., 1962). The 



1975] Chromosome Counts — Keil & Stuessy 177 



morphological and chromosomal heterogeneity within Calea 
and the absence of a recent revision of the entire genus 
suggest that a thorough modern study is much needed. 

Gimrdiola, a genus of about ten species, has been placed 
traditionally in the subtribe Melampodiinae (Hoffmann, 
1890-94). However, recent studies by the junior author 
suggest that on morphological and cytological evidence it 
belongs more properly in the Coreopsidinae (Stuessy, 
1973). Our first count of n = 12 (Fig. 8) for G. pkityphylla 
A. Gray is consistent with the recent reports of n = 12 
for both G. tulocarpus A. Gray (Grashoff et al, 1972) and 
G. mexicana H. & B. (Solbrig, Kyhos, Powell, & Raven, 
1972),- and with the base number of x = 12 for several 
other members of this subtribe.' 1 

The count of n = 10 (Fig. 9) is a first report for 
Melampodium appendicidatum Robins. In a recent revision 
of the genus ('Stuessy, 1972) this species is placed in series 
Citpulata of sect. Melampodium; three other related species 
(M. cupulatum A. Gray, M. rosei Robins., and M. tenellum 
Hook. & Arn.) also are known chromosomally as n = 10 
(Stuessy, 1971b). The addition of this new count increases 



of 37. 



species 



The first generic report for Otopappus (O. scaber S. F. 
Blake) has been published recently by Solbrig et al. (1972) 
as 7i= 16. Our first report for O. imbricatus (Sch.-Bip.) 
S. F. Blake of n = 16 (Fig. 10) confirms this chromosomal 
level for the genus. The related genera Salmea DC. and 
Notoptera Urb. (Blake, 1915) are known respectively as 
n — 18 + 2 frag. (Turner et al, 1962) and n = ca. 15 & 16 
(Turner et al., 1962; Turner & King, 1964), although very 
few taxa have been examined from each 



-The count published by Solbrig et al. was listed foe G. atriplici- 
folza A. Gray, but in the most recent published revision of the genua 
(Robinson, 1899) this epithet is regarded as 



mexicana. 



synonymous 



3 (e.g., Bidens L., Coreopsis L., Cosmos Cav., Glossorardia Cass. 
Thelrsperma Less.) 



» 



178 



Rhodora t Vo1 - 77 



thenium L., and particularly P. argentatum 



many years (cf. Hammond 
& Polhamus, 1965), including a comprehensive revision by 
Rollins (1950). Parthenium inccmum H.B.K. has been re- 
ported previously as having a polyploid series of n = 18, 
27, 36, and 45, but our new count is n = 18n & 18i (Fig. 



number 



num 



m 



mei 



genus that grow in the vicinity, such as P. 
but our voucher specimens show no morpholc 
tion of such intergradation. Alternatively, the 
figuration could indicate a hybrid between n = 18 and 
n = 36 cytotypes of the same species. The plants under 
consideration also could be apomictic, a condition that is 
known to occur in populations in the northern range of P. 
incanum (Rollins, 1950) where our material was collected. 

The count of n = 11 (Fig. 12) for Sclerocarpus spatu- 
latus Rose is consistent with previously reported numbers 
of n = 11, 12, 14, and 18 in the genus as recently defined 
by Feddema (1971). The closely related genus, Aldama 
LaLlave & Lex., is known chromosomally as n = 17 (Turner 
et al, 1962; Powell & Cuatrecasas, 1970; Feddema, 1971). 

Sigesbeckia L. of the subtribe Helianthinae is a small 
genus of less than ten species. It is worthwhile to mention 
that a close morphological resemblance exists with Tri- 
gonospermum Less. (McVaugh & Anderson, 1972; Stuessy, 



1973) 



Rumfordia 



Melampod 



thinae. Our count of n = 15 is a first report for S. agrcstis 
Poepp. & Endl. All but two other reports in the genus 
[n = 12 (Subramanyam & Kamble, 1967) and 2n = 20 
(Hsu, 1967) for S. orientalis L.] have been either n — 15 
orw= 30. Inf raspecific euploidy is known to occur in <S. 
orientalis (Mehra, Gill, Mehta, & Sidhu, 1965) and it is 
now documented for S. joj-ullensis H.B.K. by our counts of 
= 15 and 30 (Fig. 13). Only one count of n = 15 (Sol- 
brig et al., 1972) has been recorded previously for this 
species. 



n 



1975] Chromosome Counts — Keil & Stuessy 179 



Of the approximately 35 species of Simsia recognized by 
various authors (Blake, 1913, 1917, 1928; Cuatrecasas, 
1954; Robinson & Brettell, 1972), ten have been counted 
from morphologically diverse parts of the genus, and all 
counts have been n = 17. Our first counts of n = 17 (Figs. 
14 & 15) for S. eurylepis S. F. Blake and S. grayi Sch.-Bip. 
ex S. F. Blake emphasize the chromosomal uniformity 
within the genus. 

Spilanthes with approximately 60 species (Moore, 1907) 
is a taxonomically complex genus much in need of revision- 
ary attention. It appears to belong in the subtribe Galin- 
soginae rather than in the Helianthina^ as traditionally 
placed (Hoffmann, 1890-94). Chromosomally the situation 
also is complex. Even though only six species have been 
counted, four base numbers, x = 7, 12, 13, and 16, are 
present. Our first report of n = ca. 45 for S. ocymifolia 
(Lam.) A. H. Moore adds another chromosomal level to the 
already chromosomally diverse sect. "Salivaria" (= sect. 
Spilanthes) known with n = l, 12, 16, and 26. All previous 
reports for sect. Acmella (Rich.) DC. have been clearly 
based on x = 13 (only n = 13 and n = 26 counts reported). 
Our new report of n = 41 (Fig. 16) for S. phaneractis 
(Greenm.) A. H. Moore increases the chromosomal diversity 
of this section as well. 

Tricarpha is a genus of two species recently described by 
Longpre (1970). Our first count for the genus (from T. 
durangensis Longpre) of n = 8 (Fig. 17) substantiates its 
presumptive close relationship to Sabazia Cass, (n = 4, 8, 
and 16) and Selloa Kunth (n = 8) as mentioned by Long- 
pre (1970). The problem of generic delimitation in the 
subtribe Galinsoginae, involving Tricarpha, Sabazia and 

Selloa as well as Galinsoga Ruiz & Pavon, Stenocarpha 
S. F. Blake, Tridax and Jaegeria Kunth, is much in need 
of further study, despite the appearance in recent years of 
several excellent revisions (Powell, 1965; Turner, 1965; 
Torres, 1968; Longpre, 1970). Part of the difficulty in sort- 
ing out the proper affinities of all the taxa in the Galinsogi- 
nae is that previous workers have been working from the 



180 



Rhodora [Vol. 77 



perspective primarily of a single genus and not from a 
perspicacious overview of many of the genera within the 
subtribe. An added difficulty is the absence of a recent 
revision of Galinsoga (most recent treatment that of Robin- 
son, 1894), the understanding of which clearly is central to 
sorting out these generic relationships. 

Our first report for Tridax tenuifolia Rose, n = 9 (Fig. 
18), fits well with the established base number of x = 9 
for sect. THdax to which T. tenuifolia belongs (Powell, 

1965). 

Of the 14 species of Zaluzania recognized by Sharp 

(1935), six have been counted with definite numbers of 

16 and 18. Our first report, n = 17 (Fig. 19), for 



// 



mly 



new chromosomal level for the genus (a previous count of 
n = 17 ± 1 for Z. montagnae folia Sch.-Bip. was reported 

by Powell and Turner, 1963). 

helenieae. Schkuhria pinnata (Lam.) Cabrera has been 

counted before as 2n = 20 (Covas and Schnack, 1946), and 

var. virgata (LaLlave) Heiser of the same species has been 

reported as n = ca. 20 (Turner et al, 1962). Recently 

McVaugh (1972) transferred S. anthemoidea (DC.) Coult. 

var. guatemalensis (Rydb.) Heiser to S. pinnata, and our 

counts of n = 10 (Fig. 20) for this taxon are the first 

reports. A count of n = 11 (Table 1) was obtained from 



material 



However, 



■ 

our voucher specimen differs from the characters of this 
species as delimited by Heiser (1945) in having more 
numerous disc florets and large ray florets, as in S. schkuh- 
rioides (Link & Otto) Thellung in Fedde. In pappus struc- 
ture, though, our plants are much more similar to S. anthe- 
moidea than to S. schkuhrioides. This collection may repre- 
sent a previously undescribed taxon. 

senecioneae. Our first reports of n = ca. 30 for Senecio 
hartwegii Benth. and n = 22 (Fig. 21) for S. rundnatus 
Less, are consistent with counts reported previously for 
other taxa of the genus. Thirty-three species of Senecio 



1975] Chromosome Counts — Keil & Stuessy 181 



have been reported as n = 30 and six are known with n = 
22. Although the genus is based either on x = 5 (Barkley, 
1962) or x = 10 (Ornduff, Raven, Kyhos, & Kruckeberg, 
1963; Ornduff, Mosquin, Kyhos, & Raven, 1967), the diver- 
sity of haploid numbers is great, representing 30 different 
chromosomal levels from n = 5 to n = ca. 92. 

cichorieae. The small genus Pinaropappus Less, has 



from 



oseus Less., and 



the reported counts are n = 9 and 18 (Darlington & Wylie, 
1955 ; Turner et al, 1961 ; Powell & Turner, 1963 ; Powell 
& Sikes, 1970). Our present count of n = 20n & li (Fig. 
22) is a new report for this taxon. The meiotic configura- 
tions of cells in our preparation were irregular with bridges, 
lagging chromosomes, and varying numbers of univalents. 



ACKNOWLEDGMENTS 



Field work for this investigation was completed largely 
with support from NSF grant GB-30240; publication costs 
were defrayed partially by funds from NSF grant GB- 



37678. 
A. McG 



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Rhodora [ Vo1 - 77 



LITERATURE CITED 



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Bentham, G. 1873. Notes on the classification, history, and geo- 
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Jour. Bot. 53: 193-235. 



1917. New and noteworthy Compositae, chiefly Mexi- 
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Acad. Sci. 18: 25-37. 
Carano, E. 1924. Osservazioni sul meccanismo di divisione della 

cellula madre del sacco embrionale nelle piante apogame. Atti 

Reale Accad. Lincei, Rediconti CI. Sci. Fis., ser. 5, 33: 150-155. 
Cave, M. [ED.]. 1958-65. Index to Plant Chromosome Numbers, 

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Covas, G., & B. SCHNACK. 1946. Numero de cromosomas en Anto- 

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Agron. 13: 153-166. 
Cuatrecasas, J. 1954. Notas a la flora de Colombia, XIII. Revista 

Acad. Colomb. Ci. Exact. 9: 233-249. 
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Flowering Plants. Ed. 2. George Allen and Unwin, Ltd., London. 
Fagerlind, F. 1947. Macrogametophyte formation in two agamo- 



spermous Erigeron species. Acta Horti Berg. 14: 221-246. 
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posit'ae-Heliantheae). Phytologia 21: 308-314. 
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Godfrey, R. K. 1952. Pinched, section Slylimnitx, in North Ameri- 
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Grasiioff, J. L., M. W. Bierner, & D. K. Northington. 1972. 

Chromosome numbers in North and Central American Composi- 
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Hammond, B. L., & L. G. POLHAMUS. 1965. Research on Guayule 
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1975] Chromosome Counts — Keil & Stuessy 193 



Heiser, C. B., Jr. 1945. A revision of the genus Schkokria. Ann. 
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Hoffmann, O. 1890-94. Compositae, pp. 87-391. In A. Engler & 

K. Prantl, Die natiirlichen Pflanzenfamilien, vol. 4(5). Leipzig. 

Hsu, C.-C. 1967. Preliminary chromosome studies on the vascular 
plants of Taiwan (I). Taiwania 13: 117-129. 

Jackson, R. C, & R. R. Johnson. 1967, A new species of 

Machaer anther a section Psilactis. Rhodora 69: 476-480. 

Kliphuis, E., & J. H. Wiefp^ering. 1972. Chromosome numbers 
of some angiosperms from the south of France. Acta Bot. Neerl. 
21: 598-604. 

Kovanda, M. 1972. Somatic chromosome numbers for some Astera- 
ceae. Rhodora 74: 102-116. 

Larsen, K. 1953. Chromosome numbers of some European flower- 
ing plants. A preliminary note. Bot. Tidsskr. 50: 91-92. 

. 1954. Chromosome numbers of some European flower- 



ing plants. Ibid. 50: 163-174. 
Lewis, W. H. 1962. Aneusomity in aneuploid populations of Clay- 
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. 1970. Extreme instability of chromosome number in 



Claytonia virginica. Taxon 19: 180-182. 

Longpre, E. K. 1970. The systematics of the genera Sabazia, 
Selloa and Tricarpha (Compositae). Publ. Mus. Michigan State 
Univ., Biol. Ser. 4: 283-383. 

McVaugh, R. 1972. Compositarum Mexicanarum pugillus. Contr. 

Univ. Michigan Herb. 9: 359-484. 
— , & C. Anderson. 1972. North American counterparts 



of Sigrsbeckia orientalis (Compositae). Ibid. 9: 485-493. 

Mehra, P. N., B. S. Gill, J. K. Mehta, & S. S. Sidhu. 1965. Cyto- 
logical investigations on the Indian Compositae. I. North-Indian 
taxa. Caryologia 18: 35-68. 

Moore, A. H. 1907. Revision of the genus Spilanthes. Proc. Am. 
Acad. Arts 42: 521-569. 

Moore, R. J. [ed.]. 1970-72. Index to plant chromosome numbers 
for 1968-70. Regnum Veg. 68: 1-115; 77: 1-112; 84: 1-134. 

Ornduff, R. [ed.]. 1967-69. Index to plant chromosome numbers 
for 1965-67. Regnum Veg. 50: 1-128; 55: 1-126; 59: 1-129. 

, T. Mosquin, D. W. Kyhos, & P. H. Raven. 1967. 

Chromosome numbers in Compositae. VI. Senecioneae. II. Am. 
Jour. Bot. 54: 205-213. 

, P. H. Raven, D. W. Kyhos, & A. R. Kruckeberg. 1963. 



Chromosome numbers in Compositae. III. Senecioneae. Ibid. 
50: 131-139. 



194 



Rhodora [ Vo1 - 7T 



Powell, A. M. 1965. Taxonomy of Tridux (Compositae). Brit- 

tonia 17: 47-96. 

, & J. Cuatrecasas. 1970. Chromosome numbers in 

Compositae: Colombian and Venezuelan species. Ann. Missouri 

Bot. Card. 57: 374-379. 

f & S. Sikhs. 1970. Chromosome numbers of some Chi- 

huahuan desert Compositae. Southw. Naturalist 15: 175-186. 

& P L. Turner. 1963. Chromosome numbers in the 



Compositae. VII. Additional species from the southwestern 
United States and Mexico. Madrono 17: 128-140. 

"Robinson, P>. L. 1894. Notes upon the genus Galinaoga. Proc. 
Am. Acad. Arts 29: 325-327. 

. 1899. Revision of the genus Guardioht. Pull. Toiroy 



Pot. Club 26: 232-235. 

& J. M. Greenman. 1896. Revision of the Mexican and 



Central American species of the genus Galea. Proc. Am. Acad. 
Arts 32: 20-30. 

Robinson, H., & R. P. Rrettell. 1972. Studies in the Heliantheae 

(Asteraceae). II. A survey of the Mexican and Central Ameri- 
can species of Simsia. Phytologia 24: 361-377. 

Rollins, R. C. 1950. The ffuayule rubber plant and its relatives. 
Contr. Gray Herb. 172: 1-72. 

Sharp, W. M. 1935. A critical study of certain epappose genera 

of the Heliantheae-Verbesininae of the natural family Composi- 
tae. Ann. Missouri Rot. Gard. 22: 51-152. 
Solbrig, O. T. 1962. The South American species of Erigeron. 
Contr. Gray Herb. 191: 3-79. 

, L. C. Anderson, P. W. Kyhos, P. H. Raven, & L. 

Rudenbero. 1964. Chromosome numbers in Compositae V. 
Astereae II. Am. Jour. Pot. 51 : 513-519. 

, P. W. Kyhos, M. Powell, & P. H. Raven. 1972. 

— i»a ft m 



Chromosome numbers in Compositae VIII: Heliantheae. Ibid. 

59: 869-878. 
Strotiier, J. L. 1072. Chromosome studies in western North 

American Compositae. Am. Jour. Bot. 59: 242-247. 
Stukssy, T. F. 1971a. Systematic relationships in the white-rayed 

species of Melampodium (Compositae). Brittonia 23: 177-190. 
. 1971b. Chromosome numbers and phylogeny in Me- 



lampodium (Compositae). Am. Jour. Rot. 58: 732-736. 

. 1972. Revision of the tfenus Melampodium (Compo- 



sitae: Heliantheae). Rhodora 74: 1-70, 161-219. 

1973. A systematic revision of the subtribe Melampo- 



diinae (Compositae, Heliantheae). Contr. Gray Herb. 203: 65- 



80. 



1975] Chromosome Counts — Keil & Stuessy 195 



Subramanyam, K., & N. P. Kamble. 1967. In : IOPB chromosome 
number reports XII Taxon 16: 350. 

Torres, A. M. 1968. Revision of Jaegeria (Compositae-Helian- 

theae). Brittonia 20: 52-73. 

Turner, B. L. 1965. Taxonomy of Stenocarpha (Compositae-Heli- 
antheae-Galinsoginae). Southw. Naturalist 10: 238-240. 

— — , W. L. Ellison, & R. M. King. 1961. Chromosome num- 



bers in the Compositae. IV. North American species, with phy- 
letic interpretations. Am. Jour. Bot. 48: 216-223. 

, & D. Horne. 1964. Taxonomy of Maehaeranthera sect. 



Psilactis (Compositae-Astereae). Brittonia 16: 316-331. 

, & R. M. King. 1964. Chromosome numbers in the 



Compositae. VIII. Mexican and Central American species. 
Southw. Naturalist 9: 27-39. 

, M. Powell, & R. M. King. 1962. Chromosome numbers 



in the Compositae. VI. Additional Mexican and Guatemalan 
species. Rhodora 61: 251-271. 



DEPARTMENT OF BOTANY 
THE OHIO STATE UNIVERSITY 
COLUMBUS, OHIO 43210 



A VEGETAT10NAL SURVEY OF THE 
VASCULAR PLANTS OF THE KENT ISLAND GROUP, 

GRAND MANAN, NEW BRUNSWICK 1 



John.W. McCain 

In 1930, Mr. J. Sterling Rockefeller, interested in pre- 
serving the eider ducks that breed along the coast of the 
northeastern United States, purchased Kent Island, on 
which the eiders nest. In 1934, he gave the island to 
Bowdoin College as a breeding bird preserve and field 
study station. Extensive ornithological work has been done 
at this station over the years, but there have been only 
limited botanical investigations. Dr. David Potter (1936, 
1937) and John McCain (1973) compiled vascular species 
lists, and Henry A. Gleason, Jr. (1937) sampled herbaceous 
vegetation in the forested northern end of Kent Island. The 



m 



tion of the entire island and to define the plant community 
types in the non-forested areas. It is also intended to serve 
as a reference for future ornithological research on Kent 

Island. 

Kent Island lies at 66°46' E longitude and 44°35' N lati- 
tude, about fifteen miles east of the coast of Maine in the 
Bay of Fundy. It is the outermost island in the Grand 
Manan Archipelago and is politically part of Charlotte 
County, New Brunswick, Canada. Its greatest length is 



om 



mi 



•This study was supported by National Science Foundation Under- 
graduate Research Participation Grants GY-4343 in 19(58 and GY- 
(5095 in 19G9 through Bowdoin College. The author wishes to 
acknowledge the advice of Dr. E. A. Stowell of Albion College and 
Dr. E. G. Voss and Dr. W. H. Wagner of the University of Michigan, 
and the support and encouragement of Dr. Clara L. Dixon of Albion 
College and Dr. C. E. Huntington, director of the Bowdoin Science 
Station on Kent Island. This paper is published with the approval 
of the Director of the Bowdoin Science Station as Contribution 



No. 43. 



196 



1975] Kent Island — McCain 197 



All of Kent Island is now covered with water-worked 
glacial till (Meserve, 1937). The only major relief is an 
upland area with twenty-foot sand cliffs on the northern 
end of the island and two fifty-foot hills at the far .southern 
end. The northern and western ends are heavily wooded, 
and the west end is low and marshy. A 7.8 acre tidal basin 
on Kent Island is flooded twice daily by tides averaging 
14.4 feet. Kent is the largest of what local fishermen call 
the Three Islands. Fifty acre Hay Island is forested except 
for one large field on the south shore where a few now 
uninhabited houses and a weathered barn still stand. It 
can be reached on foot from Kent Island at low tide. Thirty 
acre Sheep Island is a low flat island with vegetation resem- 
bling the south end of Kent Island. It has no woody plants. 



MATERIALS AND METHODS 



summer 



ummer 



at the Bowdoin Science Station. In 1968, a general survey 
of the plant cover of the entire Three Islands group was 
made, and quantitative sampling of the vascular flora of 
the forests was completed. Aerial photographs taken in 
1965 were used to supplement the field studies in mapping 
the vegetation of the islands (Fig. 1). To determine the 
relative frequency of the forest herbs, a 100-foot grid was 
established in the wooded areas of Kent Island using a 
Brunton compass, and meter quadrats were set up at the 
intersections and at the 50-foot marks. 

The non-forested areas, identified in the 1968 survey, 
were sampled in 1969 by the line-intercept method. A non- 
forested area, as defined for the purposes of this study, is 
one in which herbs or shrubs are the dominant species and 
where trees, if present, do not form a closed canopy. Eleven 
such areas were defined. In each area, either a two- or a five- 
hundred foot transect was established along a compass line. 
Absolute and relative dominance, and relative density were 
calculated for plant species intercepted, according to the 



198 



Rhodora 



[Vol. 77 




SHEEP ISLAND 



KENT ISLAND 






****** 
****** 

****** 
' "** * 






FIELDS 

SHRUBS 

MARSHES 

GULL HILLS 
WOODED 



IOOO FT. 



FIG. I- VEGETATION MAP. KENT IS 



11)75] Kent Island — McCain 199 



method of Cottam and Curtis (1962). Absolute dominance 
is expressed in this paper as "% cover", a measure of the 
number of herbs or shrubs of a given species intercepting 
each segment of the transect line. Since several different 
species may overlap one another, the total "% cover" will 
often exceed 100%. Relative frequency was determined 
through the use of nested quadrats offset by a random 
numbers table at twenty-five foot intervals along the tran- 
sect line. The sum of relative frequency, relative density, 
and relative dominance (of woody species) is the Impor- 
tance Value (IV) of a species on a given transect. The 
sum of all species' Importance Values on each transect 
should equal 300 for shrubs, 200 for herbs. 

Soil samples were taken from the At layer at each 
sixty-five foot mark, and used to determine soil 
and water retaining capacity (WRC) by the Hilgard-cup 
method, and pH by the use of pHydrion paper. In soils 
with high organic content, the WRC values often exceed 
the dry weight of the soil itself. 

Seymour (1969) was used for the identification of the 
plant specimens with the exceptions of the two groups 
indicated in the previous paper on the Kent Island flora 
(McCain, Pike & Hodgdon, 1978). 



mois 



RESULTS 

Collections totaling 240 species and 10 varieties of vascu- 
lar plants were made on the Three Islands in 1935 by 
Dr. David Potter of Clark University, in 1968-69 by the 
present author, and in 1970 by Dr. A. R. Hodgdon and 
R. B. Pike of the University of New Hampshire. These 
collections were listed in McCain et al. (1973). Ninety-four 
of the genera in the total list (66.2%) are represented by 
one species only, as are twenty-one of the families (39% ). 
Of the fifty-four families, six (Gramineae, Cyperaceae, 
Polygonaceae, Caryophyllaceae, Rosaceae, and Compositae) 
account for 48% of the species present on the island, and 
for 100% of the species intercepted in the nesting areas of 



200 



Rhodora [Vol. 77 



• 



the islands' sizable herring gull population. The presence 
of these birds may be the main cause of the lack of floristic 
diversity. 

Eighty-five one-meter quadrats sampled in the forested 
areas showed a ground layer homogeneous in species com- 
position but widely varying in density of individuals. In 
the more open areas, or along the forest edge, Dryopteris 
spinulosa var. americana was clearly dominant and the 
total higher plant cover averaged 88.13% under spruce-fir 
and 70.56% under mountain ash and canoe birch (Be tula, 
papyrifera var. cordifolia). Where the spruce are 20-30 
feet tall, they form a very dense thicket. Here the higher 
plant cover averaged only 32.86%, with Oxalis montana 
and Maianthemum canadense dominant. In several of the 
denser spruce thickets, higher plant cover was entirely 
absent in the herb layer. The understory there suffered 
not only from a lack of light but also from a shortage of 
soil moisture. Most rain is intercepted by the dense canopy 
and most of the throughfall is caught by the deep litter 
layers before it can penetrate to the soil itself where it 
would be available for the understory plants. Mean higher 
plant cover in all eighty-five quadrats was 55.17%. Lower 
plant cover averaged 31.43% in the dense areas, 10.89% 
under the open .stands. 

The most important herb species in the wooded areas was 
Aster acuminattts (%F = 60.0). Riibus idaeus var. cana- 
densis was the most common woody species, occurring in 
28.2% of the quadrats. In openings or blowdowns, it often 
accounted for nearly 100% of the cover. Table 1 shows 
the relative frequency values for the forest floor species. 
Gleason (1937) sampled one hundred similar plots on Kent 
and Hay Islands, finding corresponding values for the spe- 
cies present. The major exception was that Oxalis montana 
was the most abundant species in his study, occurring in 
44% of his quadrats. In 1968, O. montana was present in 
24.7% of the quadrats in the total .sample, but was present 
in 71% of the quadrats in denser areas where the spruce 
was roughly 25-40 years old. This change in the presence 



1975] Kent Island — McCain 201 



of 0. montana suggests a change in the age structure of 
the tree population, with the sorrel giving way to larger 
species, often Dryopteris or the mountain aster, in the older, 
less dense woods. 



Table 1. Kent Island species occurring in more than 5 r r 



of forest area quadrats sampled, 1968. 



Species 



% frequency 



Shrubs 



Rubles idaeus L. var. canadensis Richardson 28.2 



Herbs 



Aster aciiminatus Michx, 



60.0 



Dryopteris spinulosa (0. F. Mueller) Watt. 38.8 

var. americana (Fischer) Fern. 

Rumex aceto sella L. 

Oxalis montana Raf. 

Trientalis b or calls Raf. 

Maianthemum canadense Desf. 16.4 



38.8 
24.7 
21.1 



Cor nits canadensis L. 
Poa pratensis L. 
Circaea alpina L. 



10.2 
9.4 
9.4 



Impatiens capensis Meerb. forma immacidata 8.3 

(Weath.) Fern. & Schub. 
Stellar ia media (L.) Cyrillo 8.2 

Anthoxanthum odoratum L. 7.1 

Car ex brunnescens (Pers.) Poiret 5.9 

var. sphaerostachi/a (Tuck.) Kukenthal 
Streptopus a/mplexif olius (L.) DC. 5.9 

var. americamis Schultes 



Four groups of non-forested stands can be clearly defined. 
The first includes the old fields near the middle of Kent 
Island and the main field on Hay Island. All are composed 
of turf grasses like Agrostis stolonifera and Festuca rubra 
and perennial composites such as Solidago rugosa var. 
villosa. These fields are relatively sheltered both from the 
salt spray and the weather but show the greatest influence 



202 



Rhodora [Vol. 77 



man 



Most of these areas were cleared for planting of forage 
crops, notably Phleum pratense, in the nineteenth and early 
twentieth centuries. Shrub cover is uniformly low: 1.0, 
14.40, 16.60, and 16.71% on the transects studied. Herb 
cover is the highest of the four groups, with an average 
in these fields of 386.37%. The old field transects all in- 
cluded a greater number of plant species than any of the 
other transects studied. 

The second group consists of raspberry and shrub com- 
munities. EpHobium a ngusti folium is found in association 
with Rubus idaeus var. canadensis in blowdowns and open 
areas along the north and east shores of Kent Island. Shrub 
cover values are uniformly high with a mean of 103.96%, 
but herb cover dropped to 180.37%. Each of these areas 
is the site of active gull nesting, and, as a result, has a very 
rich soil with a high organic content. 

Fresh-water and salt marsh communities along the west- 
ern shore of Kent Island and on the north shore of Hay 
Island are in the third group. Iris versicolor (Importance 
Value = 19.46), Car ex canescens var. disjuncta (IV 
23.18) and Epilobium glandulosum var. adenocaulon (IV 
= 20.65) are the most important species of these areas. 



limited 



mar 



folia and Ledum 



cum are the dominants, growing over Sphagnum moss. 
Only one small part of a formerly very extensive bog is 
left on Kent Island because of the maturing of the forests 
and man-made changes in the drainage patterns. There is 
one heath remnant, perhaps twenty feet in diameter, at the 
north edge of the bog area, the sole location found for five 
species of the flora: Calopogon tuberosus, Arethusa bnlbosa, 



aemorus and bmvetrum 



nigrum. 



In the fourth group, the South Hills of Kent Island and 
all of Sheep Island, are the main nesting areas of the 
islands' estimated 23,000 breeding pairs of herring gulls. 



U)75] Kent Island — McCain 203 



This very disturbed area supports dense populations of 
several plant species found only infrequently elsewhere on 
the island, and then only as weeds. For example, common 
ragweed (Ambrosia artemisii folia) normally loses out in 
competition to the grasses and other composites. On the 
South Hill, however, Ambrosia has an importance value of 
58.02, second only to Aster foliaceus var. arcuans (IV = 
64.62). Rumex acetosella, another weed, is similarly fav- 
ored by these disturbed conditions (IV =14.28). The 
number of species found on the South Hill is also un- 
doubtedly held down by the gulls. Only eight herb species 
and one shrub (Rubus idarus var. canadensis) appeared 
on the 500 feet of South Hill transects. By comparison, 
the four old field transects averaged 26 herb species and 
3 shrub species. 

Clearly the most frequent herbaceous species of the non- 
forest areas is the ubiquitous blue aster A. foliaceus var. 
arcuans. On all four field transects, on the north shore of 
Hay Island, and on the gull hills at the south end of Kent 
Island, it is the species with the highest importance value. 
Aster foliaceus was present in some degree in all eleven of 
the transect samples. 

The most frequent shrub species, in both forested and 
non-forest areas, is Rubus idaeus var. canadensis. Rasp- 
berry plants covered over half of the line on the four shrub 
community transects: the south field (% cover = 92), North 



Clearing (86.5'; ) Downer's Field (51%), and the Crock- 
ett's Point Field (69%). No other shrub was so clearly 
dominant in any of the areas sampled. Ribes lacustre was 
a dominant shrub (18% cover) in the Hay Island north 
shore marshes, but, where Ribes and Rubus are both pres- 
ent, the red raspberry is clearly more dominant than the 
currant. 

Soil moisture values vary from 19.5% in the North Field 
to 69.0% on the South Hill (Table 2). The South Hill also 
varies significantly in mean pH values from the rest of the 
island. The other groups of stands have mean pH values 
of 5.67, 5.77 and 5.60, while that of the South Hill is only 



204 



Rhodora t VoL 77 



4 93. WRC values are consistent within each of the four 
groups, except for a high of 629.90% on Crockett's Point 
where much poorly decomposed Sphagnum moss appeared 
in the soil profile. Soil data is summarized in Table 2. The 
extreme values for soil moisture and pH in the South Hill 
are certainly factors in the presence of a vegetation typical 
of disturbed areas. 



may 



the mechanical destruction of plants due to the perching, 
pulling and pecking of the gulls may be more harmful to 
plant survival than the chemical alteration of the soil 
caused by their droppings. 



Table 2. Soil analysis data. 

Tran- % Soil WRC 

Stand sect Moisture pH (%) 

North Field I. 19.5 6.05 121.18 



Hay Field II. 26.2 5.58 148.50 

West Field 111. 29.8 5.40 82.96 

East Field IV. 34.4 5.38 61.95 

North Clearing V. 28.0 5,80 105.55 

South Field VI. 34.7 5.70 22.64 

Downer's Field VII. 39.5 5.80 29.42 

Crockett's Point VIII. 41.0 5.80 629.90 

West shore IX. 40.0 5.70 425.44 

Hay north shore marsh X. 40.2 5.50 300.00 

South Hill XI. 69.0 4.93 196.53 



Mean Values 

Transect Group 

I-IV (Old fields) 27.48 5.67 103.50 

V-VIII (Shrubs) 35.80 5.77 *52.54 

IX-X. (Marshes) 40.10 5.60 362.72 

XL (South Hill) 69.00 4.93 196.53 

^discounting high value for transect VIII. 



1975] Kent Island — McCain 205 



The remaining botanical area of the islands is a zone just 
above the highest tide lines, favored mostly by halophytes, 
the most important among them apparently being Cakile 
edentula, Atriplex patula var. hastata and A. glabriuscula, 
Lathyrvs japonicus var. pellitus, Mertensia maritima, and 
Arenaria peploides var. robusta. There are also a few 
perennials mixed in at the upper edge of this zone, notably 
Aster foliaceus and Urtica procera. A few plants have 
ventured out onto the floor of the tidal basin where they 
are covered twice daily by the waters of high tide: Sail- 
cornia caropaea, Glaux maritima, Limonium nashii, Suaeda 
maritima and S. americana. Prominent growths of Sjxir- 
tina alterniflora and Hordeum jubatum line the edge of the 
basin. No statistical sampling has yet been done in these 
zones. 



DISCUSSION 



Three characteristics of the flora are especially worth 
noting. The first is the paucity of species. About one-half 
of the genera (56.44%) and one-fourth of the families 
(27.37%) in mainland New Brunswick are represented by 
a single species (Fowler, 1885). On Kent Island, two-thirds 
of the genera and two-fifths of the families are monotypic. 
The recentness of colonization could be a factor in the lack 
of diversity of flora, for all of the Grand Manan Island 
group was covered during the last glaciation. Thus there 
has been little time for proliferation of immigrants. The 
second possible factor is the presence of the herring gull 
colony. In work on three similar islands nearby in the Bay 
of Fundy, Hodgdon and Pike (1969) concluded that "the 
screening effect of bird populations is more apparent than 
their transport of propagules". This seemed to be true on 
Kent Island where the gulls probably prohibit by their 
activities the establishment of many potential immigrants 
and of many woody species. The continual disturbance 
caused by the birds maintains the area in early stages of 
succession. The abundance of a group of fast growing, sun 



206 



Rhodora [Vol. 77 



loving species typical of pioneer habitats, including Achil- 
lea millefolium, Rumex domesticus and R. acetosella, Urtiea 
procera, Coelopleurum lucidum and Ambrosia artimisii folio, 
indicated that these plants apparently benefited from the 

gulls' presence. 

The second notable characteristic of the flora is the pres- 
ence of a high number of boreal species for this latitude. 
Nearly three-fourths of the Kent Island .species have ranges 
that extend north and west from this area. Such species as 
Rubtfs chamaemorus, Empetrum nigrum, Vaccinium vitis- 
idaeu, Stellar ia humifusa, and Euphrasia americana reach 
their extreme southeastern limits in the Kent Island region. 
The reason for their presence here is the cool, damp climate 
of the Bay of Fundy. The warm Gulf Stream and the cold 
Labrador Current meet near the mouth of the Bay. During 
the summer the prevailing wind is from the south or south- 
west over 50% of the time. Thus the air is considerably 
warmer than the waters of the Labrador Current and of 
the Bay of Fundy (which seldom exceeds 50°F. even in 
midsummer) and the result is very frequent dense fog. 

In the ten-day period beginning 26 July 1969, for exam- 
ple, the greatest visibility was just under one mile and the 

daily low visibility was about one hundred feet. The 
fog-drip caused by the condensation of this heavy fog on 
tree limbs increases the humidity at the herb level. 

Table 4 shows pertinent weather data for the years 
1965-69. Year-round weather records for the five-year 
period 1987-1941 may be found in Cunningham's report 
(1942). 

The third important characteristic of the flora is that 
many of the new species reported in 1968-69 are weeds 
evidently introduced by man, probably after the previous 
studies were done in 1936. Thirty-one of these sixty-one 
new species were found in the field where the houses are 
on Hay Island, and on Kent Island near the wharf, along 
the main path, or in the area of the dormitory in the cen- 
tral part of the island. Conspicuous among these species 
are Tri folium pratense, Convolvulus sepium, Chenopodium 



mean 



1975] 



Kent Island 



McCain 



207 



album, Galeopsis tetrahit, Chrysanthemum leucanthemum , 
and Hieracium aurantiacum. These species are often spread 
by man, and are too conspicuous to have been missed by 
Potter in his work. 

Nearly all of Kent Island was once wooded and has been 
cut over at one time or another (Gleason, 1937). The 
northern and western ends are now wooded, with white 
spruce (Picea glauca) dominant in successional areas 
where the forest is encroaching on the fields and in blow- 
down areas. Balsam fir (Abies balsamea) is reproducing 
better in the more established woods. 



Table 3. Herbaceous and woody species cover. 







Sh 


rub cover 


Herb covei 


Stand 


Transect 




(%) 


(%) 


North Field 


I. 




16.71 


421.56 


Hay Field 


II. 




1.00 


489.50 


West Field 


III. 




16.60 


317.40 


East Field 


IV. 




14.40 


317.00 


North Clearing 


V. 




86.50 


212.50 


South Field 


VI. 




184 00 


Q3 no 



Downer's Field 
Crockett's Point Field 

West shore marsh 
Hay north shore marsh 

South Hill 



VII. 
VIII. 

IX. 
X. 

XI. 



69.00 
76 



OO 



10.00 
7.50 

16.80 



203.00 
212.97 

389.00 
156.00 

175.40 



Mean Values 
Transect group 
I-IV (Old fields) 
V-VIII (Shrubs) 
IX-X (Marshes) 
XI. (South Hill) 



12.18 
103.96 



8.75 



16.80 



386.37 
180.37 
272.50 
175.40 



208 



Rhodora t VoL 77 



Table 4. Meteorological summary of Kent Island, New 

Brunswick, Canada, 1965-1969. 
(Bowdoin Science Station Weather Records) 

Avg. Mean Avg. Mean Avg. Total Avg. Days 
Month Max.T.°F. Min. T.°F. Prec., in. Fog/Month 



June 



57.2 44.6 2.49 13.0 

July 61.8 48.8 2.44 17.8 

August 62.3 50.5 3.08 15.6 

Avg. total days with fog for the three month period : 46.4 
Mean annual precipitation (Cunningham, 1942) : 44.86" 



terms 



soil water and plant cover. Table 2 lists soil moisture con- 
tent, water retaining capacity, and pH by transects and 
Table 3 lists woody and herbaceous species cover by tran- 
sect. In the old fields, the driest areas studied, the herba- 
ceous species seemed to be encouraged. As the 



moistu 
content of the soil increased, the shrubs increased, but the 
total water the site could hold decreased as did the number 
of herbs. In the marshes the water content became high 
enough to inhibit the number of shrubs, and another group 
of herbaceous species took over. 

Finally, the highly organic soil of the South Hill had a 
high moisture content, but a lowered WRC. This is sur- 
prising since water retaining capacity usually increases 
with increased soil organic content. The increased acidity 
of these areas may have limited both the herbs and shrubs 
in variety. Where the gulls nest in areas with trees, they 
are destroying the woody plants. The south hills of Kent 
Island were formerly forested, as is evidenced by many 
fragments of decaying wood found in the soil, but the con- 
stant influence of the gulls has left only an area of twisted 
and broken snags where Rubus idaeus has now taken over. 
Many trees have been killed by the severe mechanical dam- 
age caused by the gulls. Some others have become stunted 



1975] Kent Island — McCain 209 



and have formed an unnaturally dense and tangled crown, 
thick enough and strong enough to support a man. 

In some of the most heavily disturbed areas, the grass 
pulling of the gulls keeps large areas of ground completely 
barren of all vegetation. The largest such barren area in 
1969 measured nearly fifty feet by thirty feet. The vegeta- 
tion around these barren areas is almost purely Ambrosia 
artemisii folia. This weedy species emphatically underlines 
the extreme disturbance by the gulls of this part of the 
island. Nowhere else in the Three Islands group are con- 
ditions nearly so favorable for this species. 



Literature Cited 

Cottam, G., & J. T. Curtis. 1962. Plant ecology workbook. Burgess 

Publishing Co., Minneapolis. 193 pp. 
Cunningham, R. 1942. Meteorology. Bull. Bowdoin Sci. Sta. 6: 

10-13. 
Fowler, J, 188~>. Preliminary list of the plants of New Brunswick. 

Bull. Nat. Hist. Soc. New Brunswick 4: 8-84. 
Gleason, H. A., Jr. 1937. A botanical report on Kent's Island. 

Bull. Bowdoin Sci. Sta. 3: 27-29. 
Hodgdon, A. R., & R. B. Pike. 1969. Floristic comparison of three 

bird islands in the Gulf of Maine. Rhodora 71: 510-523. 

McCain, J., R. B. Pike, & A. R. Hodgdon. 1973. The vascular 

flora of Kent Island, Grand Manan, New Brunswick. Rhodora 

75: 311-322. 
Meserve, F. W. 1937. Preliminary study of geology and geography 

of Kent's Island, 1935-1936. Bull. Bowdoin Sci. Sta. 3: 10-13. 
Potter, D. 1937. Plants found growing on Kent's Island, 1935- 

1936. Bull. Bowdoin Sci. Sta. 3: 19-26. 
Seymour, F. C. 1969. The flora of New England. Chas. E. Tuttle 

Co., Rutland. 596 pp. 

Weathered, C. A. & J. Adams. 1945. A list of the vascular plants 

of Grand Manan, Charlotte County, New Brunswick. Contrib. 
Gray Herb. Harvard Univ. No. 158. 96 pp. 



DEPARTMENT OF BIOLOGY 
ALBION COLLEGE 
ALBION, MICH. 49224 



FORM 



FROM 



Allen G. Shuey 



In 1967 several unusual plants of Thalia geniculate were 
noticed growing in a wet disturbed area caused by the 
widening of State Road 192, located ten miles west 



Melbou 



These 



become 



aquatics to the north of the road after construction on the 
road had ceased. The disturbed site was of very sandy soil 

flooded bv heavy rains. These plants 



became 



from 



Thalia by the presence of bright red petioles and petiole 

sheaths. 

For unknown reasons the plants disappeared and it was 



same 



time 



192 and west of where it was originally sited. Additional 
searching later uncovered several small populations to the 
east and west of the population found in 1972. The same 
year a stand of red-petioled Thalias was seen in a cattle 
pasture to the east of the Military Trail, northwest of the 
city of Palm Beach, Palm Beach County, Florida. This 
population has since been destroyed. 

The most striking aspect of the red form of Thalia 
geniculate is the bright rhubarb red coloration of the 
lower petiole and petiole sheaths. This coloration is carried 
farther by the presence of a red pulviniform area (usually 
greenish or orangish in the green form) at the base of the 
leaf blade, and by varying degrees of red on the joints and 
axes of the inflorescence. Red may also occur at the margin 
of the upper side of the leaf blade. 

In the green form of Thalia geniculata, no red coloration 



may 



meets 



pulviniform area at the top of the petiole. 



210 



1975] Thalia— Shuey 211 



Thalia geniculate L. f. rheumoides Shuey, forma nova 

A forma geniculata differt basibus vaginisque petiolorum 
rubris et inflorescentiae furcis, axibus, bracteisque princi- 
palibus rubris vel viridi-rubris, necnon area pulviniformi 
ad apicem petioli cremea usque rubra. 

Holotype: Florida: Brevard Co.: on south slope of water 

filled ditch, base of plant at water line, growing in full sun 

with grasses, other herbaceous plants, and the green form 

of Thalia geniculata, south side of State Road 192, 10 mi. 

west of the town of Melbourne Village, Shuey 105571 
( USF ) . 

The form name rheumoides is derived from Rheum, the 
generic name for rhubarb, and oides, Greek suffix meaning 
similar to; this was selected because the red stem sheaths 
are rhubarb red, looking like the petioles of the red vari- 
eties of rhubarb. This form of Thalia is capable of per- 
petuating itself — numerous small seedlings exhibiting red 
coloration being noted at several sites. 

The coloration of the petiole stalks and sheaths does not 
vary to any great degree from one plant to another, but 
the color of the axes of the inflorescence varies considerably. 
Some plants exhibit slight coloration only around the joints, 
bracts and bases of the inflorescence, while others tend 



mu 



mu 



light intensity. Plants growing in full sun have deeply 
colored petiole stalks and sheaths. Plants growing in the 
shade, on the other hand, have sheaths which are almost 
green, but retain the distinct red color near the margin of 
the sheath. This is lacking altogether in the green form. 

It is interesting that several plants dried for herbarium 
material retained a strong red tinge around the joints of 
the inflorescence and especially on the stem sheaths. 

I wish to extend a note of thanks to Dr. Haven C. Sweet, 
Assistant Professor of Biological Studies, Florida Techno- 
logical University, for valuable help in preparing the text 



212 



Rhodora [Vol. 77 



of this paper. Also, to Dr. Henry 0. Whittier, Assistant 
Professor of Biological Studies, Florida Technological Uni- 
versity, and to Dr. G. S. Smith (Department of Ornamental 
Horticulture, University of Florida) and Dr. D. Ward 
(Director of the Herbarium, University of Florida) for 
supplying valuable information. 



DEPARTMENT OF BIOLOGICAL SCIENCES 
FLORIDA TECHNOLOGICAL UNIVERSITY 
ORLANDO, FLORIDA 32816 



THE TAXONOMY OF TRIPOGANDRA 

(COMMELINACEAE) 1 



Wayne L. Handlos 



INTRODUCTION 

Although some specie? of commelinaceous plants — 
Tradescantia and Zebrina — are very well-known in intro- 
ductory courses in biology and botany for their use in 
demonstrating cyclosis, plasmolysis, and anther squashes, 
and while cytologists consider species of Tradescantia good 
teaching material and popular research subjects because of 
their large chromosomes, taxonomists have not agreed as 
to generic limits or relationships within the family. The 
problems of generic delimitation within the Commelinaceae 
have existed almost since the time of Linnaeus. Concepts 
have changed through time and even a careful and ob- 
servant worker such as C. Kunth modified his concepts of 
the genera during his working years as is evidenced by 
his transferral of species from one genus to another. In 
recent years the publications of Aristeguieta (1965), 
Hutchinson (1959), Matuda (1956), Moore (1960, 1963), 
Pichon (1946), Rohweder (1956), and Woodson (1942) 
illustrate the different concepts each investigator has of 
various genera, especially those American genera related 
to Tradescantia. 

Tripogandra has been included in what has been called 
the Tradescantia alliance. The United States species of 
this alliance were studied by Anderson and Woodson 
(1935) and were found to form a uniform group of species. 
Tradescantia outside of the United States contains diverse 
elements, and a study of Tripogandra was considered in 
the nature of an introduction to the whole alliance. As 



1 Based on a thesis submitted to the Graduate School, Cornell Uni- 
versity, in partial fulfillment of the requirements of the decree of 
Doctor of Philosophy. This study was supported in part by National 
Science Foundation Grant GB-6277. 



213 



214 



Rhodora [Vol. 77 



delimited by Woodson (1942) on the basis of inflorescence 
structure, Tripogandra was not well understood and con- 
tained a number of species of questionable affinity. Moore's 
(1960) investigations showed that some species had a 
modified androecium which suggested an important bio- 
logical function probably related to insect attraction, as- 
suring pollination and out-crossing. A detailed study of 
the species included in Tripogandra was thought to be 

one way of approaching the problem of generic delimita- 
tion and of determining the relationship of Tripogandra 
to its closest relatives and to the remainder of the family. 

Here I have delimited Tripogandra to include only those 
20 species which have dimorphic stamens and double cin- 
cinni not subtended by foliaceous bracts. These species are 
described in detail while the other species which have been 
included in Tripogandra at some time in the past are not 
considered congeneric and are not described in detail. 
These species and reasons for exclusion are to be found 
at the end of this paper. 



HISTORICAL REVIEW 

Tripogandra was first proposed by Rafinesque in 1837 
to contain one species, Tradescantia multiflora. Rafines- 
que's publications were not widely available and his 
taxonomic decisions were not always accepted by other 
botanists. As a result, the name Tripogandra was not 
adopted by the botanical community until much later. 
When Rafinesque published Tripogandra he also proposed 
the following as segregants from Tradescantia: Sarco peris, 
Siphonstima, Gibasis, Etheosanthes, Tripogandra (based 
on Tradescantia multiflora Jacq.) , Phyodina, Leiandra, 
Heminema (based on Tradescantia multiflora Swartz), and 
Aploleia. Rafinesque concluded his proposals with the fol- 
lowing comments: 

'These 3 last Genera [Leiandra, Heminema, Aploleia] 
lack the very essential characters of bearded Stam. that 
once was the only distinction of Tradescantia from Com- 



1975] Tripogandra — Handlos 215 



melina, but these 2 Genera are in utter confusion, as the 
above proves. Compare also my genera of Commelina. It 
is deplorable to see Botanists forcing sp. into genera, in 
spite of characters. There is not a single generic character 
common to all the above G. 22 to 32! my reform and 
revision were indispensable, and begun in 1815. . . . this 



mass 



»i 



Rafinesque was correct by present criteria in believing 
that several generically different elements were present in 
Tradescantia, but he did not solve all the problems. Both 

1 

Tripogandra and Heminema are based on the same type, 
though Rafinesque credits two different authors. Rafines- 
que may have been misled by the apparent though not real 
differences between Swartz's original description and 
Jacquin's description and illustration. Woodson (1942) 
united Hem hie ma and Tripogandra and his choice of 
Tripogandra must be followed according to Article 57 of 
the International Code of Botanical Nomenclature (1966). 

Kunth in 1843 considered the entire family and described 
sixty species of Tradescantia which he divided into several 
groups, one being "Species anomalae." This group was 
characterized by anthers of two shapes, with three longer 
and three shorter filaments. All of the species in this 
group are presently considered in the genus Tripogandra. 

Schlechtendal (1853) proposed the name Descantaria 
for the species which Kunth included in his "Species 
anomalae." Schlechtendal wrote that those species seen 
by him were distinguished by three bearded perfect sta- 
mens and three beardless imperfect stamens. No com- 
binations were made by Schlechtendal however. 

In 1866, Hasskarl used Disgrega as a generic name in 
a key with Tradescantia disgrega in parentheses probably 
indicating that this species belonged to the genus Disgrega, 
but no formal transfer of the species was made. 



om 



He divided the genus Tradescantia into three sections, one 
of which was Descantaria, characterized by three longer 



216 



Rhodora [Vol. 77 



and three shorter stamens more or less dissimilar. Clarke 
(in Donnell-Smith, 1902) described the genus Donnellia 
characterized by a three-valved capsule, locules bearing 
one seed, and based on Callisia grandi flora which was 
originally described as having two groups of dissimilar 
stamens. He was prompted to write that in his previous 
work on the family 

"no attempt is made to deal with the genera 'logically/ the 
same characters, which in the American genera (Tra- 
descantia and its allies) constitute genera, only constitute 
subgenera in Commelina and Aneilema. . . . convenience 
has been preferred to any logical system." 

Donnellia was found to be a later homonym and Rose 
(1906) proposed the name Neodonnellia for the genus. 

Clarke's work has caused concern among taxonomists 
and Anderson and Woodson (1935) note that Clarke's 

"revision of the genus Trades cantia reflects little credit 
upon the author, when compared with the critical, if ec- 
centric observation of Rafinesque nearly a half century 
previously. Not only was Clarke satisfied to ignore the 
complexity of the genus by unwarranted reducing to strict 
synonymy or ambiguous varieties most of the species pro- 
posed by his predecessors, but the literature was compli- 
cated by the publication in synonymy of unpublished names 
of other botanists, and the misinterpretation of numerous 
others. The revision of the genus in the 'Monographiae' is 
clearly a piece of chorework reflecting rather a none too 
laborious attempt at compilation than a lively interest and 

acuity." 

Bruckner (1927) chose Schlechtendal's name, Descan- 
taria, as the generic name for several of the species now 
included in Tripogandra and formally transferred those 
species. Bruckner had suggested these transfers in 1926 



com 



la 



No synonyms were given by Bruckner until 1930, when 



1975] Tripogandra — Handlos 217 



he listed Descantaria Schlechtend., Hemmema, Tripagandra 
[sic] Rafin., and Disgrega Hassk. It is not clear why 
Bruckner chose a name which lacked priority. A possible 
reason is that he had no first-hand knowledge of Rafines- 
que's publication. Bruckner's misspelling of Tripogandra 
is the same as that in Index Kewensis, fuse. 4, which may 
have been Bruckner's only acquaintance with the name. 
The American members of the Commelinaceae were 
treated by Woodson (1942) in an attempt to better delimit 
the genera. A number of transfers were made in the newly 
delimited genera, and Tripogandra was discussed at length. 
Woodson suggested new approaches to the family, stating 
that 

"the Commelinaceae always have been difficult subjects 
for herbarium study because of their deliquescent flowers. 
It is not easy to understand, therefore, why previous sys- 
tematists of the familv have focused almost their whole 
attention upon floral structure in the delimitations of sub- 
families, tribes, and genera." 

Woodson felt that stamens were too variable a character 
to use for a major subdivision of the family and pointed 
out Bruckner's inconsistency in dividing the family into 
two subfamilies and then placing Descantaria in the Hex- 
andrae and Neodonnellia in the Triandrae. I believe 
Woodson was correct in considering Descantaria and Neo- 
donnellia as congeneric. Woodson suggested that a major 
subdivision of the family could be made using inflorescence 
structure. In his concept of the tribe Commelineae, the 
ultimate units of the inflorescence are individual scorpioid 
cymes while in the Tradescantieae the basic structures "are 
paired sessile scorpioid cymes which appear as a 2-sided 
unit superficially, . . ." Woodson's concept of Tripogandra, 
which was placed in the Tradescantieae, was based pri- 
marily on inflorescence structure so it included more species 
than I have included. Woodson included species of Lep- 
torhoeo and Cuthbertia which have six similar stamens 
because these species lacked foliaceous bracts subtending 






218 Rhodora [Vol. 77 



the paired cymes. Previous authors and I have considered 
the two dissimilar whorls of stamens and their position 
as characters which delimit the most natural grouping 
of species. 

Bruckner (1930), Hutchinson (1934, 1959), Woodson 
(1942), Pichon (1946), Rohweder (1956), and Brenan 
(1966) have all dealt with the problems of generic defini- 
tion and the characters used within the family to delimit 
them. Different emphases were used by each author but 
in general a consideration of several characters was found 
necessary for more satisfactory treatments. Tripogandra 
and some of its species have been considered in recent 
years in studies of restricted scope by Macbride (1936, 
1944), Standley and Steyermark (1944, 1952), Matuda 
(1956), Moore (1960), Bacigalupo (1964, 1967), and 
Aristeguieta (1965), all of whom have been influenced 
to some degree by Woodson's work. 

The genera of the Commelinaceae have been divided 
among various groups by the different authors listed above. 
No general consensus has been reached and all schemes 
are unsatisfactory to some extent. Brenan (1966) sum- 
marized the earlier schemes of classification and has di- 
vided the genera into fifteen groups. Tripogandra is con- 
sidered in Group XI, which includes Rhoeo, Campelia, 
CaUisia, Aploleia, Tradescantia, Phiiodina, and Cymbis- 
patha. Using Brenan's criteria, Tripogandra seems more 
closely allied to the species of this group than any other. 



MORPHOLOGY 

The morphology of Tripogandra has been investigated 
to varying degrees by taxonomists who have used certain 
characters in classifying the species, but the genus has 
never been systematically investigated by a morphologist. 
Tomlinson (1966) has been a primary investigator of 
morphology and anatomy having looked at the epidermis, 
hairs and stomatal patterns of three species. Bruckner 
(1926) described in some detail the species T. glandulosa 



1975] Tripogandra — Handlos 219 



(as Tradescantia pflanzii). More recently Rohweder 
(1963a) investigated shoot development and the course of 
vascular bundles near the apex of Tripogandra pflanzii 
(— T. glandulosa). 

The following observations have been made on herbarium 
specimens and on living plants collected in Mexico or grown 
in the greenhouse. 

Habit. The plants of Tripogandra, w T hether annual or 
perennial, are succulent, weak-stemmed herbs. The plants 
may have an erect stem which is either branched or not. 
In erect, annual species — T. amplexicaulis, T. angiistifolia, 
T. guerreremis, and T. palmeri — branches may or may 
not develop from axillary buds. Plants in sunny, moist 
locations branch frequently. Crowded plants or those in 
shaded locations are commonly unbranched. Because the 
base of the plant is small and there is no secondary growth 
it is unable mechanically to support the weight of later 
growth. The base of the plant, therefore, becomes decum- 
bent and additional support and anchorage is gained by 
the production of adventitious roots at the nodes of the 
decumbent stem. The presence of an intercalary meri- 
stem at the base of each internode allows the main stem 
to remain upright through differential growth and bending 
in these areas. 

Some species, e.g. Tripogandra disgrega, T. saxicola, 
and T. purpurascens, may be either erect or trailing. This 
habit seems to be partly under genetic control ; some plants 
have sturdier stems and grow upright while others produce 
weak, flexible stems which trail over the ground. 

Perennial species such as Tripogandra montana and T. 
serrulata, commonly trail over the ground to a length of as 
much as two meters, producing adventitious roots at nearly 
every node which touches the soil. The flowering stems 
usually are erect. Branching occurs at irregular intervals. 
A vegetative branch is often produced from a node below 
the inflorescence. After growth of this axillary branch, the 
inflorescence appears to be borne in a lateral position, but 



220 



Rhodora [ Vo1 - 77 



close examination shows the vegetative shoot to be in an 
axillary position and the inflorescence to be terminal. 

Tripogandra grandiflora produces long upright shoots 
which are often supported by surrounding woody vegeta- 
tion. Sheathing bracts or cataphylls are borne on the lower 
portions of the stem. Structures transitional between the 
cataphylls and a typical vegetative leaf can be found on 

most stems. 

Root. All species of Tripogandra have fibrous roots. 
Thickened storage roots such as are found in species of 
other genera, i.e. Tradescantia, Gibasis, Setcreasea, Sepa- 
rotheca, Dichorisandra, and Commclina, have not been 
observed in any species of Tripogandra, The roots may 
be produced only at the base of the plant or they may be 
produced adventitiously from nodes along the length of 
the stem. Root initials may be differentiated early and 

as has been reported by Tomlinson (1969) 



more 



remain dormant as has been reported by Tomlinson 
for species of Tradescantia. 

Stem.. The stems of all species of Tripogandra ai 
or less succulent structures. The stems as well as most 
other organs contain a mucilaginous juice which is exuded 
when the structures are crushed or broken. A common 
feature of most species is the presence of a line or band 
of hairs extending down one side of the internode (Fig. 4). 
The hairs within this line are always uniseriate, i.e. com- 
posed of several cells joined end to end. The terminal cell 
is not enlarged or modified in shape from the cells below 
it. This line of hairs is continuous with the line of hairs 
present on the sheath of the leaf above and is always 
found on the side of the stem opposite the leaf blade of 
that sheath. The line of hairs on the internode may extend 
the full length of the internode as in T. saxicola, may 
extend for part of the length of the internode as in T. ser- 



mav 



ft 



or T. ample xicaulis. 



m 



over the surface of the stem, as in Tripogandra purpur- 
aseens subsp. au?tralis, or the stem may be glabrous as in 



1975] Tripogandra — Handlos 221 



T. grandi flora. When present, the hairs may be uniseriate 
and resemble those in the line on the internode or they 
may have an enlarged distal cell and be called capitate as 
in T. encolea. 

Leaf. The leaves of Tripogandra species are simple. The 
base is sheathing, forming a complete tube which is closely 
appressed to the internode. The leaf surface may be 
glaucous as seen in T. amplexicaulis and T. amplexans, 
but is more usually bright green and shiny. The dorsi- 
ventral blade is broad and flattened in all species except 
T. august i folia, where the lamina is C-shaped or terete in 
cross-section and in T. purpurascens where the lamina is 
complicate. The base of the blade may be variously modi- 
fied. It may be narrowed and subpetiolate as seen in some 
collections of T. amplexicaulis and T. disgrega. In some 
species, particularly on the stem just below the inflores- 
cence, the base may be amplexicaul and surround the stem 
producing a superficially perfoliate appearance as in T. 
amplexicaulis, T. amplexans, and T. encolea. In other 
species — T. diuretica, T. montana, and T. serrulata — 
the leaf base is oblique, often rounded on one side and 
cuneate on the other. 

Arrangement may be spiral as in Tripogandra guerre- 
rensis and T. disgrega or two-ranked (distichous) as most 

obvious in T. grandiflora. 

The leaves may be glabrous or variously vestite and the 
vesture may be constant or variable within a species. The 
uniseriate hairs may be distributed on the leaves in the 
following fashions : 

1. Both surfaces (abaxial and adaxial) as in some plants 
of T. disgrega and T. diuretica; 

2. Abaxially (dorsally) as in T. brasiliensis ; 

3. Adaxially (ventrally) as in T. saxicola; 

4. On the ventral surface over the midvein as in T. 

serrulata and T. montana; 

5. Adaxially in a line near the margin as in some plants 

of T. multi flora. 






222 Rhodora [Vol. 77 



Distribution of hairs is most variable in Tripogandra 



Ift 



under that species. 



The margins of blades are usually ciliate with uniseriate 



hairs except for some plants of 



ft 



and T. angustifolia. Marginal hairs may vary in shape 
from species to species and have been illustrated for 
several species by Bacigalupo (1967). The distribution 
of marginal hairs may be regular or irregular, the latter 
condition being most apparent in T. angustifolia. The 
length of the hairs varies within species. 

The orifice of the leaf sheath is villous in most species, 
although it may be glabrous in Tripogandra amplexicaulis 
and T. grandiflora. The hairs are multicellular and uni- 
seriate and intergrade with those of the leaf margin. 

Tomlinson (1969) has reported that a strand of collen- 
chyma extends along the leaf margin in Tripogandra. In 
living material this strand can be seen as a light-colored 
line and is especially obvious in T. grandiflora. 

The upper epidermis is colorless and the cells may be 
larger than any within the leaf. This phenomenon has 
been reported by Bruckner (1926) for Tripogandra pfianzii 
in his description of leaf anatomy; I have seen these en- 
larged cells in T. angustifolia and T. montana. Tomlinson 
(1966, 1969) has found this specialized epidermis in several 
genera and assumes the function to be that of water 
storage. 

The presence of silica bodies in the epidermis has been 
noted by Bruckner (1926) and Tomlinson (1966, 1969). 

The stomata have been described by Bruckner (1926) 
and Tomlinson (1966, 1969) as having two accessory cells 
adjacent to the guard cells. 

A weakly developed palisade layer may be seen in Tri- 
pogandra montana but is lacking in T. angustifolia. 

Inflorescence. The basic inflorescence unit which ap- 
pears throughout the family is the cincinnus according 

to Brenan (1966). The cincinnus as defined by Rickett 
(1955) is "a monochasium in which flowers appear alter- 
nately to right and left along one side of a sympodial axis." 



1975 ] Tripogandra — Handlos 223 



cymes 



a 



The cincinni of a plant may be arranged in various 
patterns to form more complex, compound inflorescences. 
Brenan (1966) has illustrated some of the inflorescences 
in other genera. 

The basic unit of the inflorescence in Tripogandra is 
also the cincinnus, but throughout the genus two cincinni 
occur fused together as in several other genera, e.g. Tra- 
descantia, Setcreasea, Rhoeo, and Zebrina. The type of 
inflorescence has been called the "einfache Wickel zu 
zweien" by Bruckner (1926), paired 

cinni by Woodson (1942), "Wickelpaare" by Rohweder 
(1956, 1963b), paired cincinni by Moore (1963) and 
dichotomous cymes by Mericle and Mericle (1969). 

The constant, paired nature of the cincinni in several 
genera has been described by Brenan (1966) as "a peculiar 
fusion of each pair into a characteristic bifacial unit con- 
crescent with its peduncle." Brenan has used the term 
paired cincinni" to include structures which are bifacial 
as in Setcreasea and Zebrina, as well as geminate as seen 
in Gibasis geniculates where the inflorescence is character- 
istically composed of two separate and discrete cincinni 
each on an elongate peduncle. 

I have used the term "double cincinnus" to refer to the 
bifacial, two-sided structure of the inflorescence which 
occurs in Tripogandra. The difference may be subtle but 
"double" seems to imply more in the nature of fusion than 
does "paired." Bruckner (1926) used the term "Doppel- 
wickel" which could be directly translated as double cin- 
cinnus. According to Rohweder's (1963b) translation, 
Bruckner's definition of Doppelwickel was a structure 
"composed of two cincinni arising from the same node 
and being opposite in a strict morphological sense." 
Rohweder and Brenan consider this concept erroneous. 
My use of the term double cincinnus, if it arises through 
the fusion of two separate cincinni, may conflict with 
Bruckner's concept of the origin of this structure. The 
origin of the bifacial structure has not yet been demon- 
strated. 



224 Rhodora t Vo1 - 77 



The problem of terminology is further confused by 



Rickett's (1955) 



Myosoti 



scorpioides. The structure shown corresponds to Bruck- 
ner's Doppelwickel which I believe can be seen in the 
atypical inflorescences sometimes produced in Gibasis kar- 
winskyana. I have chosen the term double cincinnus in 
lieu of a better term. 

According to the definition of a cincinnus, each flower 
is terminal and the continuing axis is always an axillary 
one. This seems to be true for Tripogandra. The pedicel 
of each flower apparently bears a bract. In the axil of 
this bract a bud develops which is terminated by a flower 
which also bears a bract on its pedicel. The small bracts 
found on the top of the peduncle of all Tripogandra species 
represent these bracts. 



seems 



Other shoots terminated by double cincinni may or may 
not be produced at lower nodes. A short shoot bearing 
several double cincinni may develop in a leaf axil, as in 
Tripogandra mvltiflora and T. montana, giving the im- 
pression that several double cincinni arise at a node. Close 
inspection shows the true nature of the situation. The 
double cincinni of T. gucrrcrensis are produced in a larger, 
much branched inflorescence so the appearance of the whole 
is that of a large panicle. 

The number of flowers produced per double cincinnus 
is variable. The cincinni of some species — Tripogandra 
montana, T. sermlata — are long-lived, producing many 
flowers over a long period of time, and the sympodial cin- 

become one or two centimeters long. On 
the other hand, T. gnerrerensis and T. amplexans, produce 



may 



ltimate 



time 



mav 



or variously vestite. The hairs may be in lines or bands 
and/or scattered. Hairs borne in lines or bands are always 
uniserate (T. serrulata) , but the scattered hairs may be 



1975] Tripogandra — Handlos 225 



either uniseriate (T. mult i flora) or capitate (T. amplexi- 
caulis) . The peduncle length is variable, probably being 
determined both genetically and environmentally. 

Flower. The flowers of all Tripogandra species follow 
the typical monocotyledonous pattern — three sepals, three 
petals, six stamens in two whorls or three stamens and 
three staminodes, and three carpels. 

The sepals, which are usually green, contain one median 
vascular bundle and are more or less boat-shaped or hooded 
near the apex. The margin is hyaline and may or may not 
be pigmented. The sepals may be glabrous or nearly so 



(Fig. 3) 



undifl 



may 



pilose with capitate hairs as in T. glandulosa and T. pur- 
purascens (Fig. 1), pilose with uniseriate hairs as in T. 
saxicola (Fig. 2), or they may have only a few hairs at 
the apex as in T. diurelica. The hairs, when present, are 
often of diagnostic value. At anthesis the sepals 
reflexed but they close when the petals deliquesce. 

The petals in all species are larger than the sepals and, 
depending on the taxon, vary in color from white to dark 
pink or magenta. Both white and pink flowers have been 
observed in Tripogandra angustifolia, T. purpurascens 

and T. sen*ulata. The petals remain open for only a few 
hours, commonly opening in the morning except for T. 
saxicola which flowers in the afternoon. All petals deli- 
quesce a few hours after opening. The cell membranes 
apparently become permeable or break down during deli- 
quescence allowing the cytoplasm to seep out of the cells 
as drops of liquid. The cell walls remain but the petal as 
a whole shrivels to a crumpled mass. The veins of the 
petals are difficult to distinguish in living material. 

The androecium is dimorphic in all species; in bud it is 
actinomorphic but at anthesis it becomes zygomorphic as 
described below. The outer whorl of the androecium is 
always opposite the sepals and is composed of fertile sta- 
mens with short filaments (Fig. 25). The filaments are 
more or less subulate or awl-shaped and may be glabrous 



226 



Rhodora t Vo1 - 77 



as in Tripogandra grandiflora (Fig. 8) or bear one to 



many multicellul 
5). The number 



(Figs. 10, 



J as a taxonomic character. In dried ma- 
terial these hairs are almost impossible to detect. In the 
living state the cells of the hairs are variable in shape so 



may 



the hairs may appear as long, unisenate struciures , 
T. palmeri (Fig. 13) or as the classical moniliform 
as in T. purpurascenx (Fig. 5). The cells of the hair 

be colorless, white, or pink. 

The distal end of the filament forms the connective 
of the anther and the connective is usually somewhat 
expanded and thickened as is easily seen in Tripogandra 
grandiflora (Figs. 8, 9). Anther sacs are borne on the 
ends of the connective (Fig. 14). The anthers are usually 
extrorse in bud, but during anthesis bending of the distal 
end of the filament causes the anthers to become introrse 
or, with less bending, pollen is exposed upward. Dehiscence 
is longitudinal in all species (Fig. 8). The pollen is spheri- 
cal to oblongoid and monosulcate. 

The structures of the inner whorl of the androecium are 
borne opposite the petals (Fig. 25) and may be either 
staminodes which produce no functional pollen or stamens 
with functional pollen. In all species the filaments of these 
stamens or staminodes are longer than the filaments of 
the outer whorl of stamens. The filaments may be glabrous 
as in Tripogandra guerrerensis (Fig. 17) and T. palmeri 
(Figs. 21, 22), or variously bearded. The hairs may be 
borne on the dorsal surface as in T. saxicola (Fig. 20), 
around the filament as in T. serrulata (Fig. 19) and T. 
montana (Fig. 31) or in two more or less discrete patches 
as in T. grandiflora (Fig. 28). In all species the hairs are 
borne more abundantly on the middle or distal end of the 
filament. The cells of the hairs are usually spherical, pro- 
ducing the typical moniliform hairs, or cylindrical, pro- 
ducing uniseriate hairs as in T. grandiflora (Fig. 28). The 
cells of one hair arc usually variable in shape and size, 
those at the base commonly being cylindrical while the 



1975 1 Tripogandra — Handlos 227 



distal ones may be either cylindrical or spheroidal depend- 
ing on the species. The cells may appear white, pigmented, 
or colorless. 

The filaments of the stamens or staminodes of the inner 
whorl are bent to varying degrees. In all species two of 
the filaments bend at the base around the filaments of two 
outer stamens so the inner stamens or staminodes become 
aligned near the third inner stamen or staminode in front 
of the upper petal (Fig. 16). In addition, all filaments 



mi 



men 



is C-shaped (Fig. 21) ; in T. serrulata (Fig. 19), T. grandi- 
flora (Fig. 28), and T. montana (Fig. 31) the bend is an 
open S-shape; while in T. guerrerensis (Fig. 17), T. 
amplexicaulis, T. amplexo,ns (Fig. 26), and T. angusti- 
folia (Fig. 30) the bend is a more pronounced S-shape. 
The filament may be more or less expanded and inflated 
distally as in T. amplexicaulis (Fig. 23) or cylindrical as 
in T. serrulata (Fig. 19). 

The connective and filament form a continuous structure 
in Tripogandra palmeri (Fig. 21) but the filament is very 
thin distally in species such as T. guerrerensis, T. amplexi- 
caulis, and T. angustifolia (Fig. 27), and the anther then 



is more or less versatile. 



may 



form 



or discoid. The anther sacs are borne on the ends of the 
connective and form a small part of the anther. Dehis- 
cence is longitudinal. 

The pollen from anthers of the outer whorl of the 
androecium may be fertile as in Tripogandra serrulata 
or modified and sterile as in T. grandiflora or T. guerre- 
rensis. Lee (1961) first described the pollen within an 
anther of tetraploid T. grandiflora. This pollen varies in 
shape from nearly spherical to oblongoid to sub-fusiform 
and absorbs stain differentially from aniline blue-lacto- 
phenol. The pollen of diploid T. grandiflora stains uni- 
formly, is more or less spherical, but is larger and has a 
more sculptured surface as compared to the pollen of the 
fertile stamens. The variation in pollen shape is greatest 



228 



Rhodora t Vo1 - 77 



in T. amplexans but within any one anther the pollen 
grains are uniform. Some collections produce spherical 



pollen which 



v _ o __ . from that of 

stamens. In other collections the pollen 



grains are oblongoid or fusiform. 

The gynoecium of the Tripogandra flower is composed 
of three fused carpels. Each carpel is supplied by three 
vascular bundles, two ventrals and one dorsal. Placenta- 
tion is axile and two orthotropous ovules are borne in each 
of the three locules (Fig. 49). A short filiform style is 
found in all species. The shape of the stigma may be 
constant or variable within a species. The stigma may 
be simple and represent the top of the style, may be slightly 
enlarged (capitellate), greatly enlarged (capitate), or 
slightly penicilliform. In some plants the stigma may be 

somewhat three-lobed. 

The pistil matures to form a loculicidal capsule which 
splits down the dorsal surface of each carpel. One to six 

be produced in each capsule. The lower ovule 
never matures in Tripogandra paJmeri, consequently a 
maximum of three seeds is produced in each capsule of 

this species. 

The orthotropous ovules develop into seeds with a dorsal 
embryotega. The embryotega represents that part of the 
integuments which lies over the embryo. In all species 



may 



emb 



um 



The micropyle is also located on the side opposite the 
hilum — the embryo, in fact, develops just below the 



mic 



m 



As has been pointed out by Bruckner (1926) 
ily and as shown by Chikkannaiah (1962, 1963, 
1964, 1965a, 1965b) from embryological studies of Com- 
melina, Murdannm, Floscopa, and Tinantia, the relation- 
ship between the micropyle, the embryo and the embryo- 
tega is a constant one. In Tripogandra pahneri (Fig. 72) 
and T. qrandiflora (Figs. 76, 77) the embryo and embryo- 



from 



of the seed are 



more 



1975] Tripogandra — Handlos 229 



surrounded and protected from mechanical damage. In 
other species of Tripogandra, the embryo is impressed 
and completely surrounded by the remainder of the seed. 
While the position of the embryotega is uniform within 
the genus it varies within the family. No detailed studies 
have been made of embryology and development of the 
seeds of Tripogandra. 

The seeds of most species of Tripogandra are trigonal 
in outline. Tripogandra amplexans has seeds which are 
variable and they may be either trigonal (Fig. 59) or 
rectangular (Fig. 58) in outline. The seeds of T. palmeri 
and T. grandffiora are unique in having revolute margins. 
Tripogandra palmeri has the margin revolute in three 



places 



din 



margins revolute (Figs. 76, 77). Seed outline is also 

■ 

dependent on the number of seeds which develop in a 
locule. The seeds become trigonal if both ovules develop 
(Fig. 76) but are more elliptical if only one develops (Fig. 
77). The single ovules usually develop into larger seeds. 

The appearance of the surface of the seed coat is of 
taxonomic value. Following the terminology of Murley 
(1951) the following categories of seed surface texture 
may be recognized among the species of Tripogandra: 

1. Reticulate: T. serrulata (Figs. 38, 39), T. montana 
(Figs. 44-46) ; 

2. Reticulate-foveate: T. multiflora (Figs. 32-35), T. 
warmingiana (Fig. 61) ; 

3. Ribbed reticulate-foveate: T. glandulosa (Figs. 42, 

43); 

4. Areolate: T. palmeri (Figs. 72, 73), T. saxicolu 

(Figs. 56, 57) ; 

5. Ribbed areolate: T. amplexans (Figs. 58-60), T. 
brasiliensis (Figs. 47, 48), T. disgrega (Figs. 52, 53), 
T. guerrerensis (Figs. 68-71), T. purpurascens (Figs. 50, 
51, 54, 55) ; 

6. Farinose: T. grandiflora (Figs. 76-78); 

7. Ribbed farinose: T. angustifolia (Figs. 74, 75). 



230 Rhodora [Vol. 77 



The seeds of Tripogandra amplexicaulis are somewhat 
variable and are generally areolate but some collections 
have seeds which are also distinctly alveolate (Figs. 64, 
65). 

Tripogandra diuretica (Figs. 40, 41) produces seeds 
which are more or less intermediate between the ribbed 
areolate and the reticulate-foveate conditions and which 
could be called ribbed areolate-foveate. 

The outline of the hilum may be punctiform (nearly 
circular) as in Tripogandra angustifolia (Fig. 75), T. 
disgrega (Fig. 53), T. diuretica (Fig. 41), T. midti flora 
(Figs. 33, 35), and T. purpurascens (Figs. 51, 55), linear 
as in T. grandiflora (Fig. 78) and T. palmeri (Fig. 73), 
or elliptical as in T. brasiliensis (Fig. 48) and T. guerre- 
rensis (Figs. 69, 71). The outline of the hilum is constant 
within a species and is useful taxonomically. 

The germination of Tradescatitia virginiana (as T. vir- 
ginica) has been described and illustrated by Gravis 
(1898) and that of Commelina virginica has been de- 
scribed by Bates (1939) but no previous studies have been 
made of Tripogandra. I have observed germination in 
detail in two species, T. amplexans and T. purpurascens. 
There are no significant visible differences in germination 
among the three genera. 

When the seeds are wet they swell slightly and within 
a few days the seedling emerges. Studies by Chikkannaiah 
(1962, 1963, 1964, 1965a) of Commelina, Murdannia and 
Floscopa, as well as those by Gravis (1898) of Tradescan- 
tia show the radicle of the embryo situated directly below 
and adjacent to the micropyle and embryotega. I have 
observed, as did Bates and Gravis, that the radicle emerges 
from the seed first. The embryotega may be pushed to 
one side or lifted free from the seed and cap the root tip 
(Fig. 24). Through further growth the remainder of the 
embryo emerges from the seed. The tip of the cotyledon 
remains in the seed, presumably as an absorbing structure, 
while the cotyledonary neck or petiole elongates further. 
The first true leaf emerges from the sheathing base of the 



1975] Tripogandra — Handlos 231 



cotyledon. By elongation of the radicle and first node, the 
seed and cotyledonary petiole may be lifted above the 
ground level. 



CYTOLOGY 

The cytological relationships of various Mexican Com- 
melinaceae as well as a review and sumary of previous 
investigations are discussed in Handlos (1970). 

i 

Tripogandra is cytologically distinct from other genera. 
Karyotypes show both telocentric and metacentric chromo- 
somes as well as larger and smaller chromosomes. The 
basic chromosome number is probably eight, though Jones 
and Jopling (1972) indicate the additional possibility of 
13. Diploid chromosome numbers range from 16 to 64. 
Two species, T. montana and T. saxicola, with haploid 
numbers of 21 may be allopolyploids based on the numbers 

8 and 13. 



BREEDING SYSTEMS AND POLLINATORS 

The genus Tripogandra is characterized by a distinc- 



tively modified androecium 



Moo 



(1960). He suggested that the arrangement of stamens 
and staminodes, in addition to the self-sterility of the clone 
studied, indicated dependence on insect pollinators and the 
necessity of out-crossing in the species. Several other com- 
melinaceous genera, including Aneilema, Cochlio sterna, 
Commelina. Tinantia. also exhibit various androecial modi- 



fications. 



been made on the ] 
roecium in the Com 



Pollination has been studied in Commelina (Breitenbach, 
1885), Tinantia (Knuth, 1906) and Tradescantia (Kerner 
von Marilaun, 1894; Sinclair, 1968). 

From a series of 166 interspecific crosses involving 
Tripogandra amplexans, T. angustif olia , T. palmeri, and 
T. pitrpurascens subsp. purpurascens made in 1966, no 
interspecific hybrids were obtained. Flowers were bagged 
to exclude insects. All species studied proved to be self- 



232 



Rhodora [Vol. 77 



fertile. Further observations of T. amplexicaulis, T. dis- 
grega, T. diuretica, T. ghndulosa, T. guerrerensis, T. 
montana, T. midtiflora, T. saxicola, and T. serrulata grow- 
ing in insect-free greenhouses show that these species are 
also self-fertile. Of the 14 species observed, only T. grandi- 
flora is self-sterile. 

In 1967, I observed natural populations of Tripogandra 
amplexans, T. amplexicmilis, T. angusti folia, T. disgrega, 
T. grandiflora, T. guerrerensis, T. montana, T. purpiir- 
ascens, and T. saxicola in Mexico for a total of 61 Vo hours. 






91 



Two hundred thirtv-seven insects were observed of which 
158 were captured. These insects are now in the collec- 
tions of the Department of Entomology, Cornell Univer- 
sity, as "Lot no. 994. 

Hymenopterous insects formed the largest category of 
visitors. There were 60 individuals in the Apidae, 48 in 
the Halictidae, 20 in the Andrenidae, 4 in the Anthophori- 
dae, 1 in the Coletidae. The bees formed the largest and 
most active group of insects and should be considered the 
most important pollinators. Bees only engaged in gather- 
ing pollen since the plants do not produce nectar. Some 
individual bees visited only the longer stamens (stami- 
nodes), others visited only the shorter, while still other 
bees crawled over all stamens and gathered pollen indis- 
criminately. 

In only one species did I observe that the position of the 
staminodes caused bees to move in a restricted fashion in 
the flower. In Tripogandra guerrerensis the petals are not 
arranged symmetrically but instead at anthesis the two 
lower petals bend away from each other and are located 
nearer the upper petal; in this way an angle of approxi- 
mately 90° is formed between either of the lateral petals 
and the upright petal, while the two lower petals are sepa- 
rated by 180°. The three staminodes arch out over the 
stamens and ovary. A bee can approach the stamens only 
by clinging upside down to the filaments of the staminodes. 
The significant position seemed to be that assumed when 
a bee was cline-ine* to the staminodes and oratheriner pollen 



1975] Tripogandra — Handlos 233 



from the anthers. The insect's abdomen then touched the 
anthers and pollen adhered to its body. It seems likely 
that pollen could be carried from flower to flower in this 
fashion and cross-pollination would thus occur. The sig- 
nificance of cross-pollination remains to be demonstrated 
in this species because these plants are also self-fertile. 
In other plants, Allard (1965) has shown that a large 
amount of variability is maintained though only a small 
percentage of outcrossing occurs. He believed that most 
individuals of a self-pollinating population would be highly 
homozygous but the recombination of genes introduced by 
a low level of outcrossing would provide sufficient new 
genotypes adapted to the microhabitats which occur in 
an area and would allow for increasing fitness of the 
species to a specific niche. Such reasoning may be applied 
to the annual and perennial self-fertile species of Tripo- 
gandra but further investigations should be conducted to 
determine the basic facts. 

Some bee species, even when collected in small numbers, 
were found to visit two or more species of Tripogandra. 
One species of Dialictns was collected from four different 
species of Tripogandra; one species of Pseudopanurgus 
and another species of Dialictns were collected from three 
Tripogandra species, while three other Dialictns species, 
one species of Evylaeus, Augochlora near smaragdina, 
Trigona fnlviv entris , T. mosqnito frontalis, and T. testacea 
orizabaensis were collected on two species of Tripogandra. 

Many of the bees collected in this study are solitary bees 
which have been reported (Linsley, 1958) to be most 
abundant in warm semi-desert regions of the world. Many 
habitats in Mexico can be described as warm and arid 



semi-desert or desert regions which may 



ma 



maj 



of solitary bees are oligolectic (Linsley, 1958) (onco- 
tropic fide Faegri and van der Pijl, 1966), i.e. utilize only 
a few related species of plants as food sources. The social 
bees, Trigona and Apis, are probably polylectic (polytropic 
fide Faegri and van der Pijl), i.e. utilize food from un- 



234 Rhodora t Vo1 - 77 



related plant sources. Trigona mexicana and T. acapulconis 
were both collected at one location in Mexico. Some indi- 
viduals were found on Tripogandra and others were caught 
on an abundant composite. 

Tripogandra grandiflora, the only demonstrated self- 
sterile species produces strongly fragrant flowers. Tripo- 
gandra ample xicaulis, T. saxicola, and T. purpurascens 
subsp. purpurascens are very faintly scented. The scents 
produced may allow bees to identify and return to a specific 
source of pollen or, in the case of T. gramdiflora, may 
allow bees to locate the plant because of its odor. 

Of 23 dipterans collected, 18 were in the family Syrphi- 
dae. Faegri and van der Pijl consider these flies to be 
irregular and not very active pollinators. In Tripogandra 
I would confirm this observation. The three dipterans in 
the family Bombyliidae may effect some pollination but 
the remaining two flies and three beetles were probably 
not visiting the flowers for pollen and should not be con- 
sidered to be imoortant oollinators. 



MEASUREMENTS 



To prevent misunderstandings and to provide for con- 
sistent results I am giving below the methods by which 
I obtained my measurements. The length of the leaf 
blade is the distance from the tip of the leaf to the top of 
the sheathing leaf base. The length of the sheath is 



mea 



from the line of attachment 



point on the orifice of the sheath; this point is always on 
the side opposite the lamina. The diameter of the leaf 
sheath reflects varying amounts of inaccuracy because 
these measurements are from dried, pressed specimens. 
If the specimen is flattened and well-pressed, the mea- 
surement will be greater than the diameter in the living 
plant; if the stem is not well-flattened, the measurement 
may be close to that in the living state or even somewhat 
smaller. In either case, the sheath seems to shrink less 
than the stem during drvinff and is a more accurate esti- 



1!,75 J Tripogandra — Handlos 235 



mate of stem diameter than a direct measurement of the 
dried stem. The length of the peduncle is the distance 
between the subtending leaf and the base of the bracts at 
the distal end of the peduncle. The length of the pedicel 

Anthesis is a definable 



measured 



period of a few hours duration and is used because it pro- 



uniform 



bias. Pedicels may elongate in fruit but a measurement 



more 



difficult to define. Measurements 
of flower parts are made on flowers at anthesis for the 
same reasons given above. The length of the filaments 
is the linear distance from the base to the connective dis- 
regarding curves and bends. The length of the anthers is 
the distance represented by the length of the anther sacs, 
while the width is taken as the distance between the anther 

Pollen viability is judged by pol- 
len stainability in aniline blue-lactophenol. The length of 
the ovary is the distance between the base of the ovary 
and the base of the style. The length of the style includes 
the stigma. The length of the capsule does not include the 
persistent style. 



same 



SYSTEMATIC ACCOUNT 

Tripogandra Rafinesque, Flora Telluriana 2:16. 1837 
('1836'). type: Tradescantia multi flora Swartz. 

Heminema Rafinesque, Flora Telluriana 2:17. 1837 
('1836'). TYPE: Tradescantia multiflora Swartz. 

Descantaria Schlechtendal, Linnaea 26:140. 1853; Bruck- 
ner, Bot. Jahr. Syst. Beiblatt 137, 61:60, 61. 1926. 
lectotype: Tradescantia multiflora Swartz. 

Disgrega Hasskarl, Flora 49:215. 1866. type: Trades- 
cantia disgrega Kunth. 

Donnellia Clarke in Donnell-Smith, Bot. Gaz. 33:261. 
1902; non Austin (1880). type: Callisia grandiflora 
Donnell-Smith. 

Neodonnellia Rose, Proc. Biol. Soc. Wash. 19:96. 1906. 
type: Callisia grandiflora Donnell-Smith. 



236 



Rhodora [ VoL 77 



Fibrous-rooted annuals and perennials, stems usually 
branching monopodially, erect or trailing, internodes cylin- 
drical, glabrous or variously vestite, often with a line of 
uniseriate hairs extending down one side. 

Leaves with a tubular sheathing base; blade narrowly 
ovate to ovate (linear in Tripogandra an g ust i folia), usually 
flat but complicate in T. purpurascens and C-shaped or 
terete in T. angustifolia, usually fleshy, glabrous or vari- 
ously vestite with uniseriate hairs, base cuneate, rounded, 
oblique, or amplexicaul, sometimes narrowed and sub- 
petiolate, apex acute, sometimes acuminate; sheath glabrous 
to pilose, orifice glabrous or more often villous with long 
uniseriate hairs, usually with a line of uniseriate hairs 
extending down the side opposite the blade and continuing 

to the internode below. 

Inflorescences composed of a single terminal or a termi- 
nal and a few axillary double cincinni, or a terminal panicle 
of double cincinni; peduncles with or without 1-2 lines of 
uniseriate hairs, otherwise glabrous to densely pilose, hairs 
uniseriate or capitate; pedicels erect or reflexed in fruit, 
glabrous to pilose, hairs uniseriate or capitate; bracts at 
the base of the pedicels small, a thin band of tissue which 
is glabrous to pilose with capitate or uniseriate hairs, 
margin entire, denticulate, with a few uniseriate hairs, 



or ciliate. 



m 



form 



glabrous to pilose, margin hyaline, entire; petals three, 
ovate, elliptic, or obovate, with acute, rounded, obtuse, or 
irregular apex, deliquescing a few hours after anthesis 
to a shapeless mass; androecium actinomorphic in bud 
becoming zygomorphic at anthesis, composed of six stamens 
or three stamens and three staminodes in two whorls, the 
outer whorl always of stamens and opposite the sepals, 
shorter, with glabrous or variously bearded filaments about 
equalling the pistil; anthers extrorse in bud, becoming 
either introrse at anthesis or horizontal and then shedding 
pollen upward toward the stigma, dehiscing longitudinally, 



1975] Tripogandra — Handlos 237 



connective usually short and inconspicuous, pollen usually 
fertile; inner whorl of stamens or staminodes opposite the 
petals, longer, filaments glabrous or variously bearded 
with uniseriate or moniliform hairs, variously curved and 
bent, two filaments bending at the base toward the third 
inner stamen-staminode around the filaments of the two 
outer intervening stamens so the inner stamens-staminodes 
are all in an erect position in front of the upper petal at 
anthesis, anthers dehiscing longitudinally, connective short 
and inconspicuous or elongate, straight or bent in a C- 
or U-shape, pollen fertile or sterile. 

Ovary globose or cylindrical, somewhat flattened on three 
sides, glabrous, trilocular, placentation axile with two 

orthotropous ovules per locule, style shorter than the ovary, 
filiform, stigma simple, capitellate, or capitate. 

Fruit a loculicidal capsule with persistent style; seeds 
usually 2 per locule (1 in Tripogandra palmeri, 1-2 in T. 
grandiflora) , usually triangular, the surface variously 
reticulate or roughened, hilum punctiform, elliptic, or 
linear, embryotega dorsal, protuberant or impressed. 



KEY TO THE SPECIES 

a. Leaves terete or C-shaped in cross section, linear in 
outline, less than 2.7 mm wide, red or green ; testa 

ribbed farinose 3. T. angusti folia. 

a. Leaves flat or complicate, never terete, mostly broader 

than 3 mm, green b. 

b. Nodes below the inflorescences bearing amplexicaul 

leaves ; leaves flat, never complicate c. 

c. Plants perennial : upper internodes pilose ; hilum 

punctiform to elliptical 7. T. encolea. 

c. Plants annual ; upper internodes glabrous or with 

a line of uniseriate hairs; hilum linear d. 

d. Petals bright pink; plants tall, to 92 cm high; 
leaves of stem below amplexicaul leaves to 
14.8 cm long, occasionally subpetiolate; pe- 
duncles pilose; seeds usually with an alveolate 
surface, convex dorsally. . 2. T. amplexicaul is. 



238 



Rhodora [Vol. 77 



d. Petals white or pale pink; plants shorter, 
to 58.5 cm high; leaves below amplexicaul 
leaves to 7.8 cm long, rounded or cuneate at 
the base; peduncles often glabrous; seeds 
never with alveolate surface, flattened dor- 
sally 1. T. amplexans. 

b. Nodes below the inflorescences without amplexicaul 
leaves or if amplexicaul then also complicate. . . e. 

e. Filaments of inner (longer) staminal whorl 
glabrous f. 

f. Sepals glabrous or nearly so; seeds two per 
locule, the upper larger, hilum linear, testa 
ribbed, areolate; inflorescence an open pani- 
cle 10. T. guerreren&is. 

f. Sepals with scattered capitate hairs or pilose; 
seeds in each locule equal in size or only one 
per locule and then the margins revolute; 
inflorescence of double cincinni variously 
arranged but never in an open panicle. . . g. 

g. Hilum linear, seeds one per locule with 
margin revolute; pedicel erect in fruit. 
15. T. palmeri. 

g. Hilum punctiform, seeds usually 2 per 
locule, lacking revolute margins; pedicel 
reflexed in fruit h. 

h. Leaf base oblique; testa reticulate but 
not ribbed; capsule obovoid, slightly 
stipitate 19. T. silvatica. 

h. Leaf base not oblique, but instead 

uneate to rounded ; testa ribbed ; 
capsule globose or obovoid, not stipi- 
tate i. 

i. Testa prominently ribbed reticu- 
late-f oveate ; longer filaments con- 
cave in upper third, to 3.3 mm long; 



<-* 



seeds 0.8-1.4 mm long 



8. T. glandvlosa. 



1975] Tripogandra — Handlos 239 



i. Testa ribbed areolate; longer fila- 
ments inflated in upper portion, to 
8.0 mm long; seeds 1.2-2.1 mm 



long. 






• •••••»••••••• 



j. Calycine hairs long, some 1.5-4.5 
mm long; leaves usually flat; 
peduncle usually glabrous; dor- 
sal seed surface convex 

5. T. disgrega. 

j. Calycine hairs short, the longest 
less than 1.0 mm long; leaves 
complicate ; peduncle variously 
glabrous, pilose, with or with- 
out lines of hairs; dorsal sur- 
face of seed flat or concave. . k. 
k. Internodes glabrous except 
for a line of uniseriate hairs 
down one side, rarely the 
upper internodes with scat- 



tered hairs. 



16a. T. purpurascens 

subsp. purpurascens. 

k. Internodes all with scattered 

capitate hairs and a line of 

uniseriate hairs down one 

side. . 16b. T. purpurascens 

subsp. australis. 
e. Filaments of inner (longer) staminal whorl 

bearded, never glabrous 1. 

1. Hairs of the filaments uniseriate, never moni- 

liform ; hilum linear, testa farinose 

9. T. grandiflora. 

1. Hairs of the filaments moniliform; hilum 

punctiform or elliptical m. 

m. Filaments of longer stamens inflated ; leaf 
base subpetiolate, oblique; testa ribbed 

areclate, hilum elliptical 

4. T. brasiliensis. 



240 



Rhodora t Vo1 - 77 



m. Filaments of longer stamens terete, never 
inflated; leaf base various; testa not 
ribbed except T. diuretica, hilum puncti- 

form (occasionally elliptical in T. saxi- 

cola and T. diuretica) n. 

n. Lamina of leaf cuneate or truncate at 

the base ; petals white o. 

o. Sepals pilose ; pedicels pilose. . . p. 

p. Pedicels erect in fruit; pe- 
duncles pilose with capitate 
hairs 11. T. kruseana. 

p. Pedicels reflexed in fruit; pe- 
duncles glabrous except for 1-2 

lines of uniseriate hairs 

17. T. saxicola. 

o. Sepals nearly glabrous, with a few 
hairs at the point of union of adja- 
cent sepals; pedicels glabrous, re- 
flexed in fruit; peduncles glabrous 
except for 1-2 lines of uniseriate 

hairs 14. T. neglecta. 

n. Lamina of leaf oblique at the base, 
rarely cuneate or rounded and then 
the petals bright pink; seeds trigonal, 
never lobed Q- 

q. Leaves subpetiolate at the base of 
the plant, to 4.4 cm long, to 1.85 
cm wide; sepal midvein pilose; 
plants annual; seeds reticulate- 
foveate ; bracts at the base of each 



pedicel with erose margi 



20. T. warming 



q. Leaves never subpetiolate, to 14.5 
cm long, to 3.5 cm wide; plants 
perennial; sepals glabrous to pi- 
lose, the midvein not distinctively 
vestite; seeds ribbed, reticulate- 



1975] - Tripogandra — Handlos 241 



foveate, or reticulate; bracts vari- 
ous r. 

r. Style nearly as long* as the 
ovary, 0.6-1.1 mm long; testa 
of seeds ribbed areolate-fove- 
ate; sepals 4.5-7.0 mm long. 

6. T. diuretica. 



•*■ 



»•••#•»«•• 



r. Style much shorter than the 
ovary, 0.15-0.6 mm long; testa 
of seeds reticulate or reticulate- 
foveate, never ribbed; sepals 

1.8-6.0 mm long s. 

s. Testa reticulate-f oveate ; pe- 
duncles pilose (rarely gla- 
brous or with 2 lines of 
hairs) ; ovary 0.5-1.0 mm 
long; sepals 1.8-4.0 mm 
long; outer anthers 0.3-0.6 
mm long. . 13. T. multiflora. 
s. Testa reticulate; peduncles 
glabrous or with 1-2 lines of 
hairs, rarely pilose, if pilose 
then ovary 1.2-1.7 mm long; 
sepals 4.0-6.0 mm long and 
outer anthers 0.8-1.8 mm 

long t. 

t. Seed surface uniformly 
brown ; sepals narrowly 

ovate, elliptical, or obo- 
vate, 4.0-6.0 mm long, 
1.6-2.8 mm wide; pe- 
duncles 0.7-12.7 cm long; 
inner filaments 4.5-7.5 
mm long, densely beard- 
ed; petals bright pink or 
magenta, 5.5-11.0 mm 
long; ovary 1.2-1.7 mm 
long; style 0.2-0.6 mm 
long. . . 12. T. montana. 



242 



Rhodora [Vol. 77 



t. Seed surface gray or 
brownish with lighter 
reticulations ; sepals 
ovate, 2.5-4.6 mm long, 
1.3-2.7 mm wide; pe- 
duncles 0.4-5.1 cm long; 
inner filaments 2.7-4.5 
mm long, variously- 
bearded; petals white or 
pink, 3.5-6.2 mm long; 
ovary 0.6-1.3 mm long; 
style 0.1-0.3 mm long. . . 
18. T. serrulata. 



1. Tripogandra amplexans Handlos, sp. nov. holotype: 
Mexico, michoacan: Km. 201.2 of Hwy. 15, 8.1 km. 



m.. 18 Sent. 1967, Handl 



(US!). 



Herba annua; caulis erectus, usque ad 58.5 cm altus, 
internodiis infra folia amplexicaulia usque ad 10.5 cm 
longis. Folia ovata, laminis usque ad 7.8 cm longis, usque 
ad 3.4 cm latis, inferis basi cuneatis, superis amplexicaulis, 
glabris, margine ciliatis, apice acutis, vaginis usque ad 
10.5 mm longis, usque ad 7.7 mm diam., glabris vel linea 
unica pilorum instructis. Injiorescentiae simplices vel pani- 
culatae ex 1-8 cincinnis duplicibus constantes, 1-5 foliis 
)lexicaulibus et 0-4 bracteis vaginantibus subtentae; 



am 



pedunculi usque ad 6.8 cm longi, glabri vel pilis paucis 
dispersis, distalibus, capitatis instructi; cincinni duplices 
alabastra, flores, vel f ructus usque ad 12 gerentes ; pedicelli 
usque ad 7.0 mm longi, pilis capitatis pilosi, erecti vel 
maturitate effusi, bracteis basi pedicellorum margine in- 
tegris, glabris vel pilis paucis, dispersis, capitatis pubes- 
centibus. Flores albi vel subrosei; sepala ovato- vel ellip- 



• ■ - 



brmia, 2.5-5.0 mm 



paucis, dispersis, capitatis, incoloribus instructa, margine 
integra et hyalina, apice acuta, rotundata vel obtusa; 



1975] Tripogandra — Handlos 243 



petala late ovata vel ovato-elliptica, 4.2-8.0 mm longa, 3.9- 
6.5 mm lata, apice acuminata, acuta, vel obtusa; stamina 
3, sepalis opposita (Fig. 25), filamentibus 1.0-2.2 mm 
longis, albis, dorsaliter medio 0-7 pilis albis, incoloribus 
vel subroseis, moniliformibus pubescentibus (Fig. 6), an- 
theris 0.3-1.0 mm longis, 0.3-1.0 mm latis, albis, polline 
albido; staminodia 3, petalis opposita (Fig. 25), filamenti- 
bus epipetalis, 2.0-6.0 mm longis, albis, glabris, sigmoideis 
et distaliter dilatis (Fig. 26), antheris luteis, 0.2-1.0 mm 
longis, 0.3-1.0 mm latis, connectivo C- vel V-formi, polline 
luteo; ovarium 0.8-1.5 mm longum, 0.7-1.2 mm diam., 
glabrum, stylo 0.3-0.8 mm longo, stigmate simplici vel 
capitellato. Capsula globosa, 2.7-3.5 mm longa, glabra, 
seminibus triangularibus raro trapezoideis (Figs. 58, 59), 
1.2-1.8 mm longis, costatis, areolatis, hilo lineari (Fig. 60). 

Chromosome number: n—16. 

Distribution and habitat: western Mexico in the states 
of Jaiisco, Michoacan, Mexico, Morelos, and Guerrero; in 
damp, rocky places in nearly neutral soil at elevations 
from 800-2200 m. 

Flowering: This species begins flowering in the rainy 
season from early August and continues until October. 
Flowers open from 8:00 AM to 9:00 AM and close between 
10:30 AM and 12:50 PM. In the field these annuals exhibit 
a great deal of variation with regard to size when flower- 
ing. This is probably due to the amount of moisture avail- 
able after germination, the effects of grazing, and the 
fertilitv of the soil. 



REPRESENTATIVE SPECIMENS 



Mexico. Jalisco: barranca SE cf Ciudad Guzman, 22 Oct. 1940, 
Moore, Jr. 158 (bh, gii). michoacan: Zitacuaro, Dist. Zitacuaro, 
1950 m., 6 Sept. 1938,. Hinlon et ah 13198 (ariz, gh, mich, mo, ny, 
us). Mexico: just N of bridge, Puente Calderon, ca. km. 135 of 
Hwy. 55, just north of Ixtapan de la Sal, 6 Aug*. 1967, Handlos 312 
(bh). morelos: railroad from Mexico City to Cuernavaca at km. 
96; region of El Parque, ca. 2200 m., 5 Oct. 1958, Hawkes, Hjerting 
& Lester 1623 (c, f). Guerrero: Manchon, Dist. Mina, 1290 m., 
13 Aug. 1936, Hinton et al 9206 (ariz, gh, ny, us). 



244 Rhodora [Vol. 77 



The range of Tripogandra amplexans is of special in- 
terest because it is wholly within the range of the more 
widespread and similar T. amplexicaulis. In many loca- 
tions the two species occur together. I am unable to dis- 
cern any differences in habitat preferences for these two 
species. 

The main reasons for considering these plants as two 
distinct species, aside from morphological differences, are 
their occurrence together with little evidence of hybridiza- 



sm 



ft base, a sma 



the slightly different flowering seasons. 

A list of distinguishing characteristics is presented here 
to summarize the differences between the species. Tripo- 
gandra amplexans is generally smaller in stature (to 58.5 
cm tall), has smaller leaves (to 7.8 cm long, to 3.4 cm 
wide) which are rounded or cuneate at 
inflorescence composed of fewer (to 8) double cincinni, a 
glabrous peduncle, flowering period from August to Octo- 
ber, fewer flowers (to 12 per double cincinnus), which 
have smaller white or pale pink petals (to 8.0 mm long), 
seeds with larger, smoother ribs, a deeply impressed 
embryotega and a flatter dorsal surface. 

Tripogandra ample xicavlis can be recognized by its 
taller stature (to 92 cm tall), its larger leaves (to 14.8 

ong, 5.0 cm wide), a narrowed, subpetiolate laminar 
base in some populations, a large paniculate inflorescence 

)osed of up to 14 double cincinni, a pilose peduncle, 
flowering period from mid-August to December, more 
flowers per inflorescence (to 18), larger 
long) , flowers bright pink, seeds with an alveolate surface, 
finer ribs, a slightly impressed embryotega and convex 
dorsal surface. 

There may be evidence of hybridization between these 
two species because at Puente Calderon in the state of 
Mexico I found one plant that was vegetatively like Tripo- 
gandra amplexans with small flowers but showing the 
bright pink color of T. amplexicaulis. Other genetic 



cm 



com 



mm 



1975] Tripogandra — Handlos 245 



mechanisms could explain the flower color difference but 
for the moment the problem has not been resolved. 

A few plants have been collected which exhibit charac- 
teristics of both species and can only arbitrarily be placed 
in either species (Figs. 29, 62, 63). Of these intermediate 
collections, five are from the known margins of the range, 
while one collected from a road cut on a steep hillside and 
another from a lava field represent populations existing 
in disturbed habitats. Further investigations should be 
conducted to determine whether these plants represent 
stabilized hybrids, populations with introgression into one 
or the other of the parental species, specialized peripheral 
populations, or whether some other explanation of their 
intermediacy is plausible. A list of the collections which 
appear intermediate follows. 



SPECIMENS EXAMINED 

Mexico. JALISCO: ca. km. 781.2 of Hwy. 15, Guadalajara to Tepic, 
ca. 2o km. NW of Ma^dalena, 1000 m., 6 Oct. 1967, Handlos 451 
(bh) ; km. 1050 of Hwy. 80, 14 km. SW of Autlan, 1120 m., 4 Oct. 
1967, Halndlos 442 (bh). Mexico: Rincon del Carmen, Dist. Temas- 
caltepec, 1340 m., 16 Sept. 1932, Hhiton 1745 (bm). MORELOS: lava 
fields near Yautepec ('Yan tepee'), 4500 ft., 22 Oct. 1902, Pringle 

8697 (bm, c, f, gh, goet, m, mexu, mo, msc, ny, ph, pom, uc, its). 

Guerrero: Rincon Viejo, 800 m., 17 Oct. 1963, Kruse 898 (mexu); 
Hwy. 95 between Ocotito and Tierra Colorado, 680 m., 21 Sept. 1967, 
Handlos 434 (bh) ; Parotas, Dist. Mina, 800 m., 12 Sept. 1936, Hinton 
et ah 9504 (ariz). 



2. Tripogandra amplexicaulis (Klotzsch ex Clarke) Wood- 
son, Ann. Missouri Bot. Card. 29:152. 1942. 

Tradescantia amplexicaulis Klotzsch ex Clarke in DC, 
Monographiae Phanerogamarum 3:304. 1881. type: 
Mexico. Chiapas, etc. Sept. 1864-70. Ghiesbreght 
887 (Lectotyoe, K; isolectotype, gh!). 

Tradescantia dilatata Clarke in DC, Monographiae 
Phanerogamarum 3:304. 1881, nom. nud. pro syn. 

Tradescantia nmbellata Pavon ex Clarke in DC, Mono- 
graphiae Phanerogamarum 3:304. 1881, nom. nud. 
pro syn. 



246 



Rhodora [Vol. 77 



Descant ana amplexicaulis (Klotzsch ex Clarke) Bruck- 
ner, Notizbl. Bot. Gart. Berlin-Dahlem 10:56. 1927. 



Plants annual; stem erect, to 92 cm tall, unbranched or 
branching profusely at the base, occasionally decumbent 
basally and rooting at the nodes; internodes below the 
inflorescence to 11.2 cm long, glabrous in the upper part 
of the plant or with a complete or partial line of uniseriate 
hairs extending down the stem from the sheath above on 
the side opposite the blade. Leaves narrowly ovate to 
ovate; blades to 14.8 cm long, to 5.0 cm wide, glabrous on 
both sides, base of variable shape, always amplexicaul 
below the inflorescence, at the base of the plant either 
cuneate, rounded, or narrowed and then appearing petio- 
late, margin ciliate, apex acuminate, or less often, acute; 
sheaths to 16.5 mm long, to 11.2 mm in diam., upper ones 
completely glabrous, lower ones with a few long uniseriate 
hairs at the orifice, and a complete or partial line of uni- 
seriate hairs extending down the side opposite the blade. 
Inflorescences terminating each stem, composed of a single 
double cincinnus or paniculate with up to 14 double cin- 
cinni, 1-6 amplexicaul leaves and 0-4 sheathing bracts, 
cincinni borne terminally and in the axils of the upper 
bracts and amplexicaul leaves; peduncles to 9.1 cm long, 
green and/or red, pilose distally, or rarely with only a few 
scattered capitate hairs with the lowest cell enlarged ; 
double cincinni with up to 13 (-18) buds, flowers and/or 
fruits; pedicels 3.0-7.3 mm long, to 1.0 mm in diam., erect 
or spreading in fruit, green or red with a green base, pilose, 
hairs capitate; bracts at the base of each pedicel pilose or 
with scattered capitate hairs, margin entire. Flowers 
bright pink; sepals ovate-cymbiform, 4.0-6.1 mm long, 1.5- 
3.0 mm wide, green w T ith a darker tip and midvein, or 
green with a red base and margin, or red, with scattered 
capitate hairs, margin entire and hyaline, apex more or 
less acute; petals ovate to broadly ovate, 6.0-13.0 mm long, 
ca. 3.4-11.5 mm wide, base cuneate, apex acuminate or 
rounded; stamens 3, opposite the sepals, filaments 1.1-2.0 



1975] Tripogandra — Handlos 247 



mm long, pink, bearing a few (5-10) pink, mo 



vs in the middle ot th< 
mm loner. 0.5-1.8 mm 



(Fig. 7), 



pink or magenta line around the more or less parallel 
anther sacs, dorsifixed, versatile, pollen whitish; stami- 
nodes 3, opposite the petals, filaments 3.5-8.0 mm long, 
slightly epipetalous, pink proximally, white distally, gla- 
brous, inflated in the distal third just below the anthers 
(Fig. 23), bent in an S-shape; anthers 0.7-2.2 mm long, 
0.7-2.2 mm wide, dorsifixed, connective yellow, V-shaped, 
with yellow, elongate more or less parallel anther sacs, 
pollen sterile, yellow; ovary 0.6-1.4 mm long, 0.7-1.3 mm 
in diam., white, glabrous, style 0.3-0.7 mm long, stigma 
simple or capitellate. Capsule 3.1-4.5 mm long, 2.5-4.0 mm 
in diam., green or brown, glabrous; seeds 2 per locule, 
triangular, 1.4-2.2 mm long, dark brown or black, testa 
areolate, sometimes alveolate (Figs. 64, 65), with ribs 
radiating from the embryotega (Fig. 66), hilum linear 
(Figs. 65, 67). 

Chromosome number: 77 = 16. 

Distribution and habitat: Sonora to Chiapas, Mexico 
and Guatemala; at elevations from (300-) 600 to 2150 m 
in thin layers of slightly acid soils which range from clay 
to sandv loam, or humus. 

Flowering : Flowering in the native habitat occurs from 
mid-August through December; most flowering collections 
seem to have been made in September and October. In 
Mexico and Ithaca, N.Y., flowers open before 8:00 AM 
and close between 11:30 AM and 12:50 PM. 

REPRESENTATIVE SPECIMENS 

Mexico, sonora: Sierra Charuco, Rio Mayo, 10 Sept. 1935, Gentry 
1706 (ariz, F, gh, mo, uc, us), chihuahua: Guayanopa Canyon, 
Sierra Madre Mts., 23 Sept. 1903, Jones (pom), sinaloa: 16.3 miles 
S.W. of El Paraiso, on road between Villa Union and El Salto, 27 
Sept. 1953, Ownbey & Ownbey 1903 (f, Mich, ny, uc, us), durango: 
La Bajada. Tamazula, 300-600 m., Nov. 1921, Ortega 4349 (us). 
nayarit: hills back of Jalisco, 11 Nov. 1925, Ferris 5981 (ds). 
Jalisco: km. 69 of Hwy. 41, N of Guadalajara, 5 Oct. 1967, 
Handlos 448 (bh). michoacan: 11 miles below Uruapan, kms. 95- 



248 



Rhodora [Vol. 77 



96, on road to Apatzing-an, 1160 m., 11 Sept. 1961, Moore, Jr. & 
Bunting 8750 (bh, MBXU, vc). Mexico : Rincon del Carmen, Dist. 
Temascaltepec, 2 Dec. 1935, Hinton et al. 8762 (ARrz, GH, mich, ny, 
us), morelos: km. 125.3 of H\\7. 95D, 6.2 km. west of bridge over 
Rio Amacuzac, ca. 1200 m., 21 Sept. 1967, Handlox 431 (bh). 
puebla : Barranca de Chochonotla, Municipio de la Union, 4 km. 
al E. de Xicotepec de Juarez, 27 Sept. 1964, Gonzalez Quintero 1674 
(MICH, MSC). Guerrero: Canon del Mano, railroad tracks north 
of Ipruala, ca. 3 km. N of El Naranjo, ca. 840 m., 13 Sept. 1967, 
Handlos All (bh). Chiapas: along Hwy. 190 in the Zinacantan 
paraje of Muctajoc, Municipio of Ixtapa, 3500 ft., 26 Oct. 1965, 
Brecdlove 13797 (ds, f, mexit). Guatemala, huehuetenango: along 
road between San Sebastian H. and San Rafael Petzal, 1900-2000 m., 
14 Aug. 1942, Steyermark 50537 (f). Guatemala: 1939, Aguilar 
374 (f). jalapa: mountains about Chahuite, northwest of Jalapa, 
about 1650 m., 16 Nov. 1940, Staindley 77472 (f). santa rosa: 
Cenaguilla, 4000 ft., Dec. 1892, Heydc & Lux 4284 (gh, m, ny, us), 
jutiapa: hills between Jutiapa and Plan de Urrutia, north of 
Jutiapa, 900-1200 m., 28 Oct. 1940, Standley 75525 (f). 

The history of the name Tripogandra amplexicaulis 
and the specimens associated with it must be considered 
to understand its present application. In the original de- 
scription, Clarke (1881) credits Klotzsch for the name 
Tradescantia ample xicaulis in the following fashion, 
"(Klotzsch ms. in herb. Berol.)". As the Berlin herbarium 
was largely destroyed during World War II it is not 
possible to examine the specimens that Klotzsch may have 
seen. However, type specimens of the Commelinaceae in 
the basement of the Berlin herbarium did not burn (Pil- 
ger, 1957) and among them is a specimen labelled "typus! 
collected by C. Ehrenberg in "Mejico" and annotated as 
Tradescantia amplexicaulis by C. B. Clarke in his own 
hand. This may be the specimen Klotzsch would have 
designated as the type for his name. Klotzsch never pub- 
lished his description so this specimen has no standing 
under the current rules. Clarke does not mention the speci- 
men in his description of the species. The Ehrenberg 
specimen represents Tripogandra amplexans. Clarke listed 
eight collections of Tradescantia amplexicaidis (Schaffner 
108, 138; Botteri 531, 892; Ghiesbreght 887; Salvin; 
Savage; Hoffman) which are syntypes because no holo- 



yy 



1975 3 Tripogandra — Handlos 249 



type was designated. Additionally he cited Tradescantia 
umbellata Pa von as a synonym and I have seen one speci- 
men so labelled from the British Museum and annotated 
by Clarke as T. dilatata, his manuscript nrme for T. 

amplexicaulis. 

The Pavon specimen (T. umbellata), Schatfner 108, and 
Botteri 581 represent Tripogandra purpurascens. Botteri 
892 and Ghiesbreght 887 represent the taxon presently 
under consideration and the remaining specimens have 
not been seen. The original description given by Clarke 
does not overwhelmingly refer to any one species. To 
preserve current usage of the name T. amplexicaulis I 
therefore designate Ghiesbreght 887 (K) as the lectotype 
for T. amplexicaulis. The following phrases from Clarke's 
description are taken as referring to T. amplexicaulis 
sensu stricto and distinguishing it from T. purpurascens. 
"Pedunculis quasi paniculam efformantibus, . . . Folia 
. . . acuminata, . . . vaginae ore glabratae; folia summa 



fere ad vaginas 



tempore patenti- 



erecta." The remainder of Clarke's original description 
could apply to both species. I do not understand his state- 
ment, "Ovarium apice vix aut minute pubescens." I have 
never seen hairs on the ovary of any specimens of Tripo- 
gandra. 

In the past Tripogandra amplexicavHs and T. amplexans 
have been considered the same species. I prefer to separate 
them on the basis of a number of morphological characters, 
a difference in flowering season and the fact that I have 
found both species growing side by side with little or no 
evidence of hybridization as discussed under T. amplexans. 

3. Tripogandra angustifolia (Robinson) Woodson, Ann. 

Missouri Bot. Gard. 29:152. 1942. 

■ 

Tradescantia angustifolia Robinson, Proc. Amer. Acad. 
Arts 27:185. 1893. type: Mexico, san luis potosi: 
Las Canoas, 14 Aug. 1891, Pringle 3902. (Holo- 
type, gh!; isotypes. b!, bm !, br!, e!, goet!, m!, mo!. 

msc!, ny!, ph!, uc!). 



250 



Rhodora [Vol. 77 



Descantaria angu&tifolia (Robinson) Bruckner, Notizbl. 
Bot. Gart. Berlin-Dahlem 10:56. 1927. 

Illustrations: Matuda, Anales Inst. Biol. Nac. Mexico 
26:368. 1956 ('1955'). 

Plants annual; stem erect, to 24 cm tall, unbranched 
or branched; internodes to 5.0 cm long, green or red, 
glabrous except for a line of uniseriate hairs extending 
straight down the side opposite the leaf blade from the 
sheath above. Leaves linear; blades to 7.1 cm long, 2.7 mm 
wide, terete or C-shaped in cross section, glabrous dorsally 
and ventrally, margin usually ciliate in the proximal por- 
tion or occasionally to the tip, acute at the apex; sheaths 
to 4.1 mm long, to 4.0 mm in diam., with a few long uni- 
seriate hairs at the orifice, otherwise glabrous except for 
a line of uniseriate hairs extending down the side opposite 
the lamina. Inflorescences composed of 1-4 double cincinni 
borne terminally and in the axils of the upper leaves; 
peduncles to 7.5 cm long, green or red, glabrous or some- 
times with 1-2 lines of uniseriate hairs extending down one 
side; buds, flowers and/or fruits 2-10 per double cincinnus; 
pedicels to 8.0 mm long, to 0.6 mm in diam. at anthesis, 
green or red, glabrous, erect in fruit; bracts at the base 
of each pedicel glabrous, margin entire, erose, or some- 
times ciliate, lowest 1-2 bracts sometimes leaf-like. Flowers 
pink, occasionally white; sepals elliptic- or ovate-cymbi- 
form, to 4.5 mm long, to 2.0 mm wide, green, green with 
a darker tip, or red, glabrous, margin entire, hyaline, apex 
acute; petals broadly ovate, to 5.0 mm long, to 4.0 mm 
wide, cuneate basally, apex acute; stamens 3, opposite the 
sepals, filaments to 1.5 mm long, white or pink, nearly 
glabrous except for a few minute hairs, anthers 0.3-0.8 mm 
long, 0.4-0.7 mm wide, pink, connective bent in the middle 
and U-shaped with the anther sacs parallel (Fig. 14), 
pollen white; staminodes 3, opposite the petals, filaments 



mm long, pink and/ 



mm 



bent in an S-shape distally (Fig. 30), anthers 0.7-1.1 
long, 0.7-1.1 mm wide, versatile, connective yellow, promi 



li)7f) ] Tripogandra — Handlos 251 



nent, C-, U- or V-shaped, with the yellow anther sacs di- 
vergent (Fig. 27), pollen white, cream, or yellow, sterile; 
ovary 0.5-1.0 mm long, 0.4-1.0 mm in diam., green, gla- 
brous, style 0.3-0.7 mm long, stigma simple or capitellate. 
Capsule globose, 2.0-3.0 mm long, 1.5-2.4 mm in diam., 
brown, glabrous; seeds 2 per locule, triangular, 0.9-1.3 mm 
long, brown or gray, testa ribbed farinose (Fig. 74), hilum 
punctiform (Fig. 75), embryotega impressed. 

Chromosome number: n~S. 

Distribution and habitat: central and southern Mexico 
and western Guatemala from the states of San Luis Potosi, 
west to Guerrero and south to Chiapas in Mexico and the 



Guatemal 



limest 



Flowering: In the native habitat, flowering occurs in 
August through October. In a uniform environment, plants 
collected at different locations flower at different times, 
probably indicating physiological variation in response to 
differing environmental factors in the various localities. 
Flowers open between 7:00 AM and 8:15 AM — earlier 
in sunny locations and later in shady locations. Closing 
occurs between 11:00 and 11:30 AM. 

REPRESENTATIVE SPECIMENS 

Mexico, san luis potosi: Pozo de Azuna, 15 km. al E. de Gua- 
dalacazar, 1450 m., 26 Sept. 1955, Rzedowski 6695 (mexu, mich). 
hidalgo: between Jacala and Barranca Seca via Hilo Juanico, Dist. 
Jacala, 1400-1600 m., 30 Oct. 1946, Moore, Jr. 1800 (bit, gh). Vera- 
cruz: Banos del Carrizal, Aug. 1912, Purpus 6174 (f, gh, mo, ny, 
UC, us). MEXICO: rounded hill with crater in the center about 0.5 
km. N. of village of Tonatico, 23 Aug. 1965, Handlos 169 (bh). 
morelos: near Yautepec, Aug. 1903, Rose & Painter 6575 (gh). 
puebla: km. 298.3 of Hwy. 190, ca. 12.8 km. south of Acatlan, 1340 

m., 11 Sept. 1967, Handlos 406 (bh). Guerrero: km. 231.6-7 of Hwy. 
95, 37.7 km. north of Chilpancingo, 30 Aug. 1967, Handlos 388 (bh). 
oaxaca: Cerro de San Antonio de la Cal, 1700 m., 18 Aug. 1907, 

Conzatti 1995 (f, mich). Chiapas: km. 1062.5 of Hwy. 190, 4 km. 
south of Berriozabal, 930 m., 18 Aug. 1967, Handlos 364 (bh). 
Guatemala, huehuetenango: dry slopes between San Ildefonso 
Ixtahuacan and Cuilco, 1350-1600 m., 16 Aug. 1942, Steyermark 
50694 (F, MO, US). 



252 



Rhodora [ Vo1 - 77 



Tripogandra angustifolia is unique for its narrow leaves 
which are nearly terete or C-shaped in cross-section. These 
leaves are probably a special adaptation which allows the 
plants to store water between rains and grow in very dry 
areas. Small plants can survive and produce seeds with 
as few as three or four leaves and two to four flowers in 
a single terminal inflorescense. To illustrate how densely 
plants of T. angustifolia grow, in 1967 I arbitrarily se- 
lected an area of 100 square cm, ten cm to a side. I care- 
fully removed each plant and discovered there were exactly 
240 plants in that area! Each plant was surviving on an 
average of 0.416 square cm of space. 

This species seems to be the only one possessing very 
short hairs on the staminal filaments. These obscure hairs 
may indicate that the ancestor of Tripogandra angustifolia 
had a bearded filament and that as an adaptation for water 
conservation these structures were reduced but not com- 



pletely lost. 



til 



their triangular outline and the conspicuously ribbed sides. 
Maintenance of this species in cultivation has proved a 
problem because seed germination has been very low. 
Field-collected seeds have produced one or two or no seed- 
lings at all where seeds nearly covered the surface of a 
pot of soil. This contrasts with conditions in Mexico where 
plants grow in very dense stands, implying a high per- 
centage of seed germination. 

Tripogandra angustifolia is morphologically very uni- 
form throughout its range which extends for some 1270 
km from north to south. The variations which occur — 
white or pink flowers, red or green stems or leaves — occur 
within a population and do not distinguish one population 

another. One or two leaf-like bracts occur rarely 



om 



may 



similar 



Tradescantia 



which has large leaf-like bracts. There are slight varia- 
tions in intensity of flower color and petal shape in plants 



1975] Tripogandra — Handlos 253 



from different populations, but the significance of these 
variations is unknown. The general lack of morphological 
variation may be due to the specialized and uniform habitat 
which this species inhabits — the thin layer of alkaline 
soil found in cracks or on ledges of limestone rock through- 
out southern Mexico and parts of Guatemala — where it 
seems to be specially adapted to survive. As soil depth 
increases, other species of plants seem to be better adapted 
to compete and T. an g list i folia is crowded out. 

4. Tripogandra brasiliensis Handlos, sp. nov. HOLOTYPE: 
Brazil, maranhao: "Ilha de Balsas 91 region, between 
the Balsas and Parnaiba Rivers. Ca. 6 km. north of 
main house of Fazenda "Morros", ca. 30 km. south of 
Loreto, ca. 300 m., 30 April 1962, Eiten & Eiten 4458 

(NY!). 

Herba annua (?) ; caulis erectus, usque ad 52.5 cm altus, 
internodiis usque ad 13.4 cm longis, linea unica pilorum 
instructis, aliter glabris vel pilis dispersis pubescentibus. 
Folia angusto-ovata, petiolata, laminis usque ad 8.7 cm 
longis, usque ad 2.1 cm latis, apice acuminatis, basi obliquis, 
margine ciliatis, dorsaliter pilosis vel pilis dispersis in- 
structis, ventraliter glabris praeter lineam unicam pilorum 
secus costam, vaginis usque ad 6.0 mm longis, usque ad 
3.6 mm diam., orificio villosis, linea unica pilorum in- 
structis, aliter glabris vel pilis dispersis praeditis. Inflo- 
rescentiae terminates et in axillis foliorum summorum, 
ex 1-2 cincinnis duplicibus constantantes ; peduncidi usque 
ad 3.9 cm longi, pilis paucis dispersis pubescentes; pedicelli 
usque ad 3.0 mm longi, glabri, bracteis basi pedicellorum 
glabris, margine integris. Flore s albi ; sepala ovato-cymbi- 
formia, usque ad 4.2 mm longa, usque ad 2.7 mm lata, 
glabra vel 1-2 pilis instructa, margine hyalina, apice 
obtusa; petala non visa; stamina 6 in verticillis duobus, 
3 sepalis opposita filamentibus brevis, usque ad 1.5 mm 
longis, glabris, antheris usque ad 0.7 mm longis, usque 
ad 0.6 mm latis, 3 petalis opposita filamentibus longioribus, 
usque ad 3.9 mm longis, dilatatis et barbatis distaliter, 



254 



Rhodora [Vol. 77 



mm long-is, usque ad 0.9 mm 



connectivo elongato; ovarium usque ad 1.5 mm longum, 



usque ad 1.0 mm diam., glabrum 



stigmate 



mm 
mm 



mm 



laribus, usque ad 2.0 mm longis, costatis, areolatis (Fig. 
47), hilo elliptico (Fig. 48). 

Vernacular name: baixao do cipo, fide Eiten & Eiten. 

Distribution: known from only three locations in 
Maranhao and Minas Gerais, Brazil. The habitat of this 
species, according to Eiten and Eiten, was tall forest along 
a gully in a disturbed area with a pronounced dry season 
and intermittent streams. 



SPECIMENS EXAMINED 

Brazil, maranhao: 30 April 1962, Eiten & Eiten 4458 (ny). 
minas gerais: Lagoa Santa, Warming (f p.p., us p.p.) ; La#oa 
Grande, Belo Horizonte, Mar. 1935, Cocheau (r). 

This species is recognizable from its probable annual 
habit, petiolate leaves, peduncle with scattered uniseriate 
hairs, an inflated filament on the longer stamens, and the 
large ribbed areolate seeds with elliptical hilum. While 
the collections known to me are few, the species is un- 
doubtedly distinct from other South American species of 

Tripogandra. 

5. Tripogandra disgrega (Kunth) Woodson, Ann. Missouri 

Bot. Gard. 29:152. 1942. 

Tradescantia disgrega Kunth, Enumeratio Plantarum 
4:97. 1843. type: Mexico. Serro Colorado, Aug. 
1828, Schiede 974 (816) (Lectotype, b!; isolectotype, 

HAL!). 

Tradescantia ehrenbergiana Klotzsch ex Clarke in DC, 
Monographiae Phanerogamarum 3:305. 1881, nom. 
nud. pro syn. 

Disgrega mexicana Hasskarl ex Clarke in DC, Mono- 
graphiae Phanerogamarum 3:305. 1881, nom. nud. 
pro syn. 



1975 ] Tripogandra — Handlos 255 



Descantaria disgrega (Kunth) Bruckner, Notizbl. Bot. 
Gart. Berlin-Dahlem 10:57. 1927. 

Tradescantia disgrega forma glandulosa Standley & 
Steyermark, Field Mus. Nat. Hist, Bot. 'Ser. 23:36. 
1944. holotype : Guatemala, zacapa : Sierra de las 
Minas, along trail between Rio Hondo and summit 
of mountain at Finca Alexandria, 1000-1500 m., 11 
Oct. 1939, Steyermark 29751 (f!) ; photograph (f!). 

Tradescantia disgrega forma pubescens Standley & 
Steyermark, Field Mus. Nat. Hist., Bot. Ser. 23:37. 

1944. holotype: Guatemala. Guatemala: Borra.ja, 

1085 m., Oct. 1928, Morales R. 1106 (f!) ; photo- 
graph (f!). 

Tripogandra disgrega forma glandulosa (Standley & 
Steyermark) Standley & Steyermark, Fieldiana : 
Bot. 24(3) :37. 1952. 

Tripogandra disgrega forma pubescens (Standley & 
Steyermark) Standley & Steyermark, Fieldiana: 
Bot. 24(3) :37. 1952. 

Plants annual, rooting at lower nodes; stem erect, to 
55 cm tall, branched or unbranched, some plants with the 
lowermost portion decumbent; internodes to 12.5 cm long, 
pilose or glabrous except for a line of uniseriate hairs ex- 
tending down the side of the stem, continuous with the 
line on the sheath above. Leaves narrowly ovate to ovate; 
blades to 9.8 cm long, 3.8 cm wide, cuneate, rounded, or 
subpetiolate at the base, glabrous to pilose on both sides, 
the uppermost leaf usually with fewer or no hairs, margin 
ciliate, apex acute or acuminate; sheaths to 15.0 mm long, 
4.3 mm in diam., villous at the orifice, a line of uniseriate 
hairs extending down the side opposite the blade, other- 
wise glabrous to pilose. Inflorescences composed of 1-5 
double cincinni borne terminally and in the axils of the 
upper leaves; peduncles to 9.5 (-15) cm long, glabrous or 
with a few capitate hairs near the distal end; double cin- 
cinni with up to 25 buds, flowers and/or fruits; pedicels 
to 7.0 mm long, glabrous proximally but pilose distally, 



256 



Rhodora [Vol. 77 



reflexed in fruit ; bracts at the base of each pedicel pilose 
or with scattered capitate hairs, margin entire. Flowers 
pink; sepals ovate- or elliptic-cymbiform, to 6.0 mm long, 
3.0 mm wide, green or with red at the tip and/or base, 
pilose or with scattered capitate hairs 1.5-4.5 mm long, 
margin hyaline, apex more or less acute; petals broadly 
ovate, to 8.0 mm long, to 7.0 mm wide, base cuneate, apex 
acuminate; stamens 6, in two whorls, the outer shorter, 
filaments to 2.0 mm long with a few (ca. 10) moniliform 
hairs borne on the middle of the dorsal side, anthers 0.7-1.6 
mm long, 0.5-1.4 mm wide, dorsifixed, versatile, with 
anther sacs slightly spreading (not parallel) ; stamens of 
the inner whorl longer, filaments to 8.0 mm long, glabrous, 
with a U- or open S-shaped bend and inflated in the upper 
half, anthers to 1.5 mm long, to 1.6 mm wide, basifixed, 
connective conspicuous, anther sacs either divergent or 
becoming parallel and adjacent through bending of the 
connective in a U-shape; ovary to 1.1 mm long, to 1.3 mm 
in diam., glabrous, style 0.4-0.7 mm long, stigma simple 
or capitellate (minutely penicilliform) . Capsule globose, 
to 3.5 mm long, to 3.5 mm in diam., green or brown at 
maturity, glabrous ; seeds 2 per locule, triangular or rarely 
elliptical, 1.2-1.7 mm long, brown to black, testa areolate, 
with ribs (sometimes obscure) radiating from the em- 
bryotega (Fig. 52), hilum punctiform (Fig. 53). 

Vernacular name : yerba del polio fide Hinton. 

Distribution and habitat: from Jalisco to Chiapas, 
Mexico, Guatemala, Honduras and El Salvador; in moist 
areas alonjr streams or in pine forests, occasionally a weed 
in cornfields. 

Flowering: This species flow r ers from August to Novem- 
ber in different parts of Mexico, and from October to 
February in Guatemala, The flowers open before 8:45 AM 
and begin closing about 11 :25 AM. 



REPRESENTATIVE SPECIMENS 



Mexico. Jalisco: Sierra del Tigre, 3 miles south of Mazamitla, 
2100-2200 m., 22 Sept. 1952, McVaugh 13167 (Mini), hidalgo: 
vicinity of Molan&o on Lolotla road, Municipality Molan^o, District 



1975] Tripogandra — Handlos 257 



Molango, 1600 m., 9 Nov. 1946, Moore, Jr. 1984 (bh). veracruz: 
Orizaba, Botteri 326 (Gh). michoacan: roadsides in pine zone about 
48 kms. from Patzcuaro on road to Tacambaro, ca. 6-7000 ft, 2 Sept. 
1948, Moore, Jr. & Wood, Jr. 4847 (bh), Mexico: Ypericones, Dist. 
Temascaltepec, 23 Nov. 1935, Hinton et al. 8718 (GH, MICH, mo, ny). 
morelos: in pine forest, km. 61, off Cuernavaca-Mexico City High- 
way, 2 Oct. 1943, Lundell & Lundell 12496 (mich, ny, uc). oaxaca: 
15 km. al S. de Sola de Vega, sobre la carretera a Puerto Escondido, 
1800 m., 30 Sept. 1965, Rzedowski 21296 (mich). Chiapas: sitio in 
San Cristobal las Casas, Municipio of San Cristobal las Casas, 7100 
ft., 20 Sept. 1965, Breedlove 12310 (ds). Guatemala, san marcos: 
barrancos 6 miles south and west of town of Tajumulco, north- 
western slopes of Volcan Tajumulco, 2300-2800 m., 26 Feb. 1940, 
Steyermark 36621 (f). chimaltenango: above Las Calderas, 1800- 
2100 m., 15 Dec. 1938, Standley 60014 (f). sacatepequez : Ciudad 
Vieja, Nov. 1914, Tejada 301 (us). Guatemala: near San Juan 
Sacatepequez, about 1800 m., 8 Dec. 1938, Standley 59258 (f). santa 
rosa: Santa Rosa, 3000 ft., Nov. 1892, Heyde & Lux 4285 (gh, ny, 
us). Honduras. MORAZAN: colinas cultivadas de maiz de la Monta- 
nita, 1500 m., 18 Nov. 1948, Molina R. 1610 (GH, us), el paraiso: 
slopes above Yuscaran, Montserrat, 1500 m., 25 Nov. 1958, Hawkes, 
Hjerting & Lester 2061 (c, k). El Salvador, santa ana: Hda. Los 
Planes nordl. Metapan, 1800 m., 29 Oct. 1950, Rohweder 707 (f). 
morazan : eastern edge, finca of General J. T. Calderon, Montes de 
Caca^uatique, ca. 1340 m., 28 Dec. 1941, Tucker 646 (bh p.p., F p.p., 

MICH p.p., MO, NY p.p., PH p.p., UC p.p., US p.p.). 



Schlechtendal (1831) described but did not name a plant 
of Tripogandra disgrega from a specimen (Schiede 974 
(815)) housed at Halle. Kunth later named the species 
and his description was drawn from a duplicate specimen 
of Schiede's housed in Berlin "(Descr. juxta specimen 
Schiede.)". In all respects the description and specimen 
are in accord. I have seen two sheets of this collection 
and because there is no evidence that Kunth saw the sheet 
at Halle, I designate the Berlin specimen as the lectotype, 
therefore making Schlechtendal's material at Halle an iso- 
lectotype. 

Kunth's original description cites another specimen, 
Berlandier 948. There is such a specimen in the Berlin 
herbarium but it is not annotated by Kunth so perhaps 
he did not see this particular sheet. This collection is a 



258 



Rhodora [Vol. 77 



specimen of Tripogandra purpurascens. These two species 

lar and, as the Berlandier specimens are widely 



are sim 



distributed, the name T. disgrega has been applied con- 
sistently to two different species. As Kunth drew his 
description from the Schiede specimen, no name other than 
T. disgrega can be applied to this species. There seems 
to be no other course if one believes there are two species 



name 



uui i>vy vw** vax*^ v **^ — - — — , .- - - & ~- — & «. * 

to the species represented by Berlandier 948. 

Tripogandra disgrega has some variable morphological 
characteristics but I am unable to see any trends or pat- 
tern in the variation. The species is rare and has not been 
collected often. The vesture of the leaves varies consider- 
ably but not in any consistent way. The leaves may be 
glabrous on both sides or may have a varying amount of 
hair ranging from scattered to pilose. When hairs are 
found on the leaves they are also present on the inter- 
nodes and on the leaf sheaths. Tripogandra disgrega fo: 



ma 



extreme 



simi 



There does seem to be some correlation between the 
presence of hairs on the leaves and a narrowing at the 
base of the blade producing a subpetiolate appearance. 
More of the pilose-leaved plants have a narrowed blade 
than do the glabrous-leaved plants, but the significance, if 
any, of this correlation is not evident. 

Tripogandra disgrega and T. purpurascens are very 

most respects and on this basis I would con- 
very closely related. One difference separates 
all specimens and a second difference separates most. 
Tripogandra disgrega has long hairs on the sepals, the 
longest from 1.5-4.5 mm long, while the hairs on the sepals 
of T. purpurascens are shorter and vary from 0.2-1.0 mm 
long on dried specimens. Most plants of T. disgrega have 
a glabrous peduncle, but in three of 31 collections there 
were scattered capitate hairs on the distal end. Tripo- 
gandra purpurascens is more variable, plants may have 
glabrous peduncles or any single plant may have both 



em 



1 975] Tripogandra — Handlos 259 



glabrous and variously vestite peduncles, ranging from 
scattered hairs to pilose and/or with one or two lines of 
hairs. 

The following differences between these species generally 
hold but there are exceptions which may be due partly to 
environmental influences, may be due also to genetic effects, 
or could be the result of undetected hybridization. Tripo- 
gandra disgrega usually has flat open leaves which may be 
subpetiolate, the plants grow in shady habitats, the petals 
are acuminate distally, the dorsal surface of the seeds is 
rounded or convex, the seeds are often obscurely ribbed, 
and in any given location the peak of flowering seems to 
occur later in each season than the peak of flowering for 
T. purpurascens. In contrast, T. purpurascens has compli- 
cate (folded or canaliculate) leaves which are usually 
rounded at the base and never narrowed and petiolate, 
the plants grow in open sunny locations, the petals are 
irregularly indented or acuminate at the apex, the seeds 
are prominently ribbed and the dorsal surface is flat or 
even concave with a deeply impressed embryotega, and 
flowering seems to occur earlier in each season in a given 
location than the peak of flowering for T. disgrega. 



6. Tripogandra diuretica (Martius) Handlos, comb. nov. 

Tradescantia diuretica Martius in Spix & Martius, 
Reise in Brasilien 281. 1823. HOLOTYPE: Brazil. 
Min. Ge. et S. Pauli, Martius (m!). 

Tradescantia commelina Vellozo, Florae Fluminensis 
140; 3:154. 1829 (1825). holotype: apparently 
lost. 

Tradescantia diuretica ft foliis vaginisque magis glabris 
Schultes in Schultes & Schultes, Systema Vegeta- 
bilium 7:1163. 1830. HOLOTYPE: Brazil. Martius 
(m!). 

Tradescantia gaudichaudiana Kunth, Enumeratio Plan- 
tarum 4:93. 1843. holotype: Brazil. Rio Janeiro, 
1832, Gaudichaud 125 (b!). 



260 



Rhodora t Vo1 - 77 



Tradescantia mollis Kunth, Enumeratio Plantarum 
4:95. 1843. TYPE: Brazil. 1836, Sella 565 (Holo- 
type, b!; isotype, b!). 

Tradescantia sello wiana Kunth, Enumeratio Plantarum 
4:93. 1843. type: Brazil. Vittoria-Bahia, 1836, 
Sello 1006 (Holotype, b!; isotype, b!). 

Tradescantia diuretica « mollis (Kunth) Seubert in 
Martius, Flora Brasiliensis 3(1) :251. 1855. 

Tradescantia diuretica glabriusada Schultes ex Seu- 
bert in Martius, Flora Brasiliensis 3(1) :251. 1855. 
holotype: Brazil. Yrino, Sello 1309 (b!). 

Tradescantia elongata & diuretica (Martius) Clarke in 
DC, Monographiae Phanerogamarum 3:303. 1881. 

Descantarm diuretica Schlechtendal fide Hasskarl ex 

Clarke in DC, M 

304. 1881, nom. nud. pro syn. 



urn 3: 



Tripogandra elongata forma diuretica (Martius) 
Standley & Steyermark, Fieldiana: Bot. 24(3) :38. 
1952. 

Illustrations: Bacigalupo, Darwiniana 14:407, 410. 
1967; Figs. 4, 5k, 1, as Tripogandra elongata. 

Plants perennial, the base decumbent, rooting at the 
nodes; stems to 11.5 dm long, flowering stems erect, rarely 
branched; internodes to 12.1 cm long, with a complete or 
partial line of uniseriate hairs extending down the side 
from the sheath above, otherwise glabrous or rarely com- 
pletely pilose or pilose distally. Leaves narrowly ovate to 
ovate ; blades to 14.2 cm long, to 2.6 cm wide, base oblique, 
rounded on one side, cuneate on the other, dorsal and 
ventral side glabrous to pilose, mid vein with or without 
a partial or complete line of uniseriate hairs ventrally, 
margin ciliate; sheaths to 20.0 mm long, to 9.5 mm in 
diam., villous at the orifice, with a line of uniseriate hairs 
extending down the side opposite the blade, otherwise 
glabrous to pilose. Inflorescences of 1-10 double cincinni 
borne terminally and in the upper 1-4 leaf axils ; peduncles 
to 8.9 cm long, usually glabrous, rarely pilose or with a 



1975] Tripogandra — Handlos 261 



few scattered uniseriate hairs; buds, flowers and/or fruits 
to 17 per double cincinnus; pedicels 3.5-9.0 mm long, 
glabrous, reflexed in fruit; bracts at the base of each 
pedicel glabrous or rarely pilose or with a few scattered 
uniseriate hairs, margin entire, with a few uniseriate 
hairs, or ciliate. Flowers pink; sepals narrowly ovate to 
ovate, cymbiform, 4.5-7.0 mm long, 1.4-3.3 mm wide, 
glabrous or with a few uniseriate hairs at the apex, rarely 
pilose, margin hyaline, apex acute; petals more or less 
elliptic, to 10.0 mm long, to 6.5 mm wide, base cuneate, 
apex acute to rounded ; stamens 3, opposite the sepals, fila- 
ments 1.2-2.3 mm long, glabrous or with a few (2-3) hairs, 



mm loner. 0.6-1.1 mm 



more 



4.0-6.3 mm long, bearded in the upper third or fourth 
with moniliform hairs, distal end bent in an S-shape, an- 
thers 0.7-1.4 mm long, 0.8-1.5 mm wide, basifixed, with 
parallel, yellow anther sacs, pollen sterile; ovary 0.7-1.4 
mm loner. 0.7-1.0 mm in diam., glabrous, style 0.6-1.1 mm 



tigm 



mm 



mm in diam 



mm 



light gray, testa ribbed, areolate-foveate 
elliptic or punctiform (Fig. 41). 



um 



Chromosome number: 2n— 64 (A. Sparrow, pers. 

comm.). 

Vernacular name: trepueraval, jupirava tupice fide 

Martius; trepoerava fide Peckolt, Brazil. 

Distribution and habitat: southern Brazil, Bolivia, 
Paraguay, Argentina, and Uruguay; commonly in low wet 
places near sea level to 1600 (-2200) m. This species in 
Brazil seems adapted to moist swampy locations and is not 
found in limestone outcrop areas as are many of the species 
in Mexico (Father Reitz, verb. comm.). 

Flowering: Flowering occurs from December to July. 



REPRESENTATIVE SPECIMENS 



Bolivia, la paz: San Bartolome, near Calisaya, Basin of Rio Bopi, 
province of S. Yun^as, 750-900 m., 1-22 July 1939, Krukoff 1054.". 



262 



Rhodora [Vol. 77 



(GH, k). cochabamba: vie. Cochabamba, 1891, Bang 1282 (bm, 
E, F, G, GH, NY, us). Paraguay. Mbuveva, Apr. 1931, Jorgenaen 4355 
(us). Brazil, pernambuco : Pesqueira, 20 Jan. 1943, Vellozo 561 

(r). matto grosso: Smith 818 (r). minas gerais: Vigosa, road 

from Canella to Repressa, Buraco Canella, 730 m., 7 Jan. 1930, 

Mexia 5487 (bh, bm, f, gh, la, mich, mo, ny, ph, uc, us), rio de 
Janeiro: Theresopolis, na Serra dos Orgaos, Jan. 1955, Vidal V-120 
(r). FEDERAL district: Botanical Garden and vicinity, Nov. 1915, 
Curran 7 (us). SAO paulo: Parque do Estado de Sao Paulo, 26 
Mar. 1931, Hoehne 27392 (b, ny, sp). Parana : Jaguariahyva, 28 
Mar. 1916, Dusen 18037 (f, mo, ny, us), santa catarina: Mina 
Velha, Garuva, S. Francisco do Sul, 10 m., 26 Mar. 1958, Reitz & 
Klein 6618 (ny, uc, us), rio GRANDE no sul: Torres, Municipio 
de Torres, Feb. 1939, Vidal (r). Uruguay. Montevideo: Miguelete, 
10-20 m., May 1925, [ Herter 149 (b, bh, f, gh, m, mich, mo, msc, 
ny, U, UC, Wis). Argentina. MISSIONES: Dep. San Pedro, Loc. 
Avellaneda, 8 Apr. 1949, Schwindt 1535 (br, ny, us), buenos aires: 
Partido de Delta, Paraya Mini, 18 May 1950, Cabrera 10635 (us). 

The plants which I have included in Tripogandra diure- 
tica generally have been considered a form of T. elongata 
by taxonomists. The differences in the seeds, flowers, ratio 
of style length to ovary length, leaf vesture when present, 
and habitat are of a greater magnitude than those which 
serve to distinguish forms. Furthermore, I consider T. 
elongata sensu stricto to be composed of sterile material 
which I include in T. serrulata. The problem is further 
discussed under that species. 

Variation in leaf vesture is seen among the collections 
of Tripogandra diuretica, but does not seem to follow any 
discernible geographic pattern. 

Herbarium material of this species is unusual because 

a high percentage of specimens possesses open, dried flowers. 

This is related to the fact that flowers on plants growing 

in the greenhouse remain open all day and are longer lived 
than those of other species. 



7. Tripogandra encolea (Diels) Macbride, Revista Univ. 

(Cuzco) 33(87) :142. 1944. 

Tradeacantia encolea Diels in Urban, Bot. Jahr. Syst. 
37:381. 1906. type: Peru, cajamarca: San Pablo, 



1075] Tripogandra — Handlos 263 



2200-2400 m., 1906, Weberbauer 3855 (Holotype, b!; 
isotype, g!). 
Descantaria encolea (Diels) Bruckner, Notizbl. Bot. 
Gart. Berlin-Dahlem 10:56. 1927. 

"Tripogandra encolea (Diels) Rohweder", Abh. Aus- 
landsk., Reihe C, Naturwiss. 18:156. 1956. 

Plants trailing, decumbent, stems branching and rooting 
at the nodes; flowering stems upright, to 65 cm tall, 0.1- 
0.4 cm in diam. when dry, green; internodes 1.5-9.5 cm 
long, always with a line of uniseriate hairs extending in a 
line laterally down the internode from the sheath above, 
otherwise glabrous basally and pilose distally in the erect 
part of the plant. Leaves narrowly ovate; blade 1.9-7.4 
long, 0.9-2.6 cm wide, reduced to sheathing bracts near the 
top of the flowering stem, tapering or cordate at the base 
on the lower part of the stem, usually amplexicaul at the 
base below the inflorescence, glabrous ventrally, glabrous 
dorsally or sometimes pilose near the sheath, hairs uni- 



cm 



cm 



cm in diam 



wise glabrous except for a line of uniseriate hairs laterally 
opposite the leaf blade; sheathing bracts glabrous, pilose, 
or with scattered capitate hairs. Inflorescences composed 
of 1-4 double cincinni borne terminally and in the axils of 
the upper leaves; peduncles 1.0-7.0 cm long, ca. 1 mm in 
diam. when dried, glabrous or pilose at the base, densely 
pilose near the apex, hairs capitate, brown or colorless; 
double cincinni with 7-18 buds, flowers and/or fruits; 
pedicels of flowers near anthesis 0.3-0.7 cm long, pilose 
with capitate hairs; bracts at the base of each pedicel 
glabrous or pilose with capitate hairs. Flowers white or 
pink; sepals ovate-cymbiform, 4.5-6.2 
wide, pilose with brown or colorless capitate hairs, apex 
obtuse, margin entire; petals obovate, ca. 4-5.8 mm long, 
ca. 2.5-4.5 mm wide, tapering at the base, apex obtuse; 
stamens of outer whorl 3, shorter, filaments before anthesis 
1.3-2.0 mm long, glabrous, anthers 0.9-1.5 mm long, 0.7- 



mm 



264 Rhodora [Vol. 77 



1.0 mm wide, white, dorsifixed, versatile, with parallel 
anther sacs; stamens of inner whorl longer, filaments 3.0- 
3.5 mm long, densely bearded with moniliform hairs in the 
distal portion, anthers 0.8-1.0 mm long, 0.7-1.1 mm wide, 
basifixed, versatile, with yellow anther sacs parallel or 
diverging slightly, longer than the connective; ovary 0.9- 
2.0 mm long, 0.8-1.2 mm in diam., glabrous, style 0.3-0.7 
mm long, stigma simple or capitellate. Capsule brown, 
glabrous, ca. 3 mm long, 2.5 mm in diam.; seeds 6, tri- 
angular, 1.3-1.5 mm long, dark brown; hilum punctiform 
to elliptical. 

Distribution and habitat : Peru to Bolivia ; in moist rocky 
soil. 



SPECIMENS EXAMINED 

Peru. CAJAMarca: San Pablo, 2200-2400 m., 1906, Wehcrbaver 
3855 (b, g). la libertad: Sammne-Casmiche, Prov. Otuzco, 1800 m., 
21 May 1952, Lopez M. 0844 (us), lima: San Mateo, Prov. Huaro- 
chiri, 3200 m., 24 Mar. 1952, Hutchison 815 [grown and collected 
Univ. of Calif.-Berkeley, 19 Oct. 1957, G.R. Newcomb] (bh, f, g, 
gh, UC). Matucana, ca. 8000 ft., 12 April-3 May 1922, Macbride & 
Featherstone 350 (G, us) ; Prov. Cajabomba, Banos de Churin, 9000 
ft., 7 Feb. 194G, Laudcmer 5414 (k). Bolivia, cochabamba: Ayo- 
paya, Sailapata, 2700 m., Cardenas 3029a (us). Rio de Guinllabamba, 
10 June 1876, Andre 3f>08 (k, ny). 



Diels' original description of this species reported the 
length of leaf sheaths to be 3-4 cm, the peduncles to be 
1.5-2 cm, and petals to be 8-9 mm. My measurements of 
the type specimens show leaf sheaths 0.7-2.9 cm long, 
peduncles 1.0-4.5 cm long, and petals 5.8 mm. If additional 
information was available to Diels he does not mention 
that fact. 

An examination of all the specimens available of this 
rather rare species reveals that there are minor morpho- 
logical differences among them which is not surprising if 
one considers the limited amount of material available 
and the ecological diversity of the mountains which this 
species inhabits. 



1975] Tripogandra — Handlos 265 



8. Tripogandra glandulosa (Seubert) Rohweder, Abh. 

Auslandsk., Reihe C, Naturwiss. 18:156. 1956. 
Tradescantia glandulosa Seubert in Martius, Flora 

Brasiliensis 3(1) :253. 1855. holotype: Brazil. 

Parana: Rio Negro, 7 Mar. 1823, SeUo 995 (b!). 
Tradescantia radlata Clarke in Chodat & Hassler, Bull. 

Herb. Boissier 3:245. 1903. HOLOTYPE: Paraguay. 

amambay: in regione cursus superioris fluminis Apa, 

Feb. 1901-2, Hassler 8493 (g!). 
Tradescantia pflanzii Bruckner, Bot. Jahr. Syst. 61(1) : 

13. 1927 ('1926'), nom. mid, 
Descantaria glandulosa (Seubert) Bruckner, Notizbl. 

Bot. Gart. Berlin-Dahlem 10:56. 1927. 
Descantaria pflanzii Bruckner, Notizbl. Bot. Gart. Ber- 
lin-Dahlem 10:57. 1927. holotype: cult, in hort. 

Berol. (Bl). 
Descantaria radiata (Clarke) Bruckner, Notizbl. Bot. 

Gart. Berlin-Dahlem 10:56. 1927. 
Tripogandra pflanzii (Bruckner) Rohweder, Abh. Aus- 
landsk., Reihe C, Naturwiss. 18:156. 1956. 
Tripogandra radiata (Clarke) Bacigalupo, Darwiniana 

13:90. 1964. 

Illustrations: Bacigalupo, Darwiniana 13:92, 93. 1964 
(Figs. 1, 2) ; 14:404, 410. 1967 (Figs. 3, 5). 

Plants perennial, rooting at nodes of decumbent stems; 
flowering stems upright, to 39 cm tall, branching from 
decumbent stems ; internodes to 9.5 cm long, glabrous or 
with a partial line of uniseriate hairs extending down the 
stem from the sheath above. Leaves narrowly ovate to 
ovate ; blades to 6.7 cm long, to 2.3 cm wide, glabrous, base 
cuneate on the lower part of the plant, rounded on the 
upper portion of the plant, margin ciliate, apex acute; 
sheaths to 11.0 mm long, to 8.6 mm in diam., a few long 
hairs at the orifice, a line of uniseriate hairs extending 
down the side of the sheath opposite the blade, otherwise 
glabrous. Inflorescences of 1-10 (-23) double cincinni borne 
terminally and in 1-4 of the upper leaf axils; peduncles 



266 



Rhodora t Vo1 - 77 



cm 



seriate hairs, or glabrous proximally and pilose distally, 
hairs capitate; double cincinni with up to 11 buds, flowers 
and/or fruits; pedicels 2.3-6.0 mm long, recurved in fruit, 
pilose with capitate hairs; bracts subtending each pedicel 
glabrous to pilose with capitate hairs, margin denticulate, 
entire, or ciliate. Flowers white or pink; sepals narrowly 
ovate, 3.0-4.2 mm long, 1.2-1.9 mm wide, pilose, hairs capi- 
tate, margin hyaline, apex acute; petals ca. 3.5-5.0 mm 
long ; stamens 6, in two whorls, the outer shorter, filaments 
to 1.5 mm long, with moniliform hairs borne on the mid- 
dorsal portion, anthers 0.6-0.9 mm long, 0.4-0.8 mm wide, 
basifixed, anther sacs parallel ; stamens of the inner whorl 
longer, filaments to 3.3 mm long, glabrous, ligulate, con- 
cave in the upper third of their length, anthers 0.4-1.1 mm 
long, 0.4-0.7 mm wide, basifixed, anther sacs parallel; 
ovary 0.5-1.0 mm long, 0.4-0.9 mm in diam., glabrous, 



mm long, stigma simple or cap 
.7 mm loner, to 3.1 mm in diam 



Capsule 



brown ; seeds 2 per locule, rounded-triangular, 0.8-1.4 mm 
long, light gray to brown, testa reticulate-foveate with 
small prominent ribs radiating from the punctiform hilum 

(Figs. 42, 43). 

Chromosome number; w— 8. 

Distribution and habitat: southern Bolivia and Brazil, 
Paraguay, Uruguay, and northeastern Argentina; in moist 

places and along streams. 

Flowering: Flowering of this species seems to occur 

between September and March. 

REPRESENTATIVE SPECIMENS 

Brazil. Parana: Sete Quedas, Mun. Guaira, 11 Dec. 1965, Hatsch- 
bach, Lindeman & Haas 13339 (us). Paraguay, amambay: zwischen 
Rio Apa und Rio Aquidaban, 1908-9, Fiebri« 4487 (bm, G, k, l). 
SAN pedro : Alto Paraguay, Primavera, 13 Oct. 1955, Woolxton 596 
(C, NY). GUARIA: Villarrica, Cerro Peladu(?), Dec. 1930, JorgensCn 
4112 (l)S, F, mo, ny, PH, US). Argentina. Formosa: Colonia Clorinda, 
150 m., 30 Dec. 1926, Venturi 9163 (us), catamarca: road 38 from 
Tucuman to Catamarca, Cuesta de Totoval, km. 1384.7, Dept. Paclin, 

900 m., 24 Mar. I960, Hawkes, Hjerting £ Rahm 3989 (c). tucuman: 



1975] Tripogandra — Handlos 267 



El Duraguito, Dept. Capital, 550 m., 14 Jan. 1922, Venturi 1672 
(f p.p., us, US p.p.). CHACO: Margarita Belen, Dept. Resistencia, 
9 Dec. 1945, Aguilar 563 (g). SANTA fe: Lonteri, Estancia Romaz- 
zola, 1 Feb. 1936, J oh 1185 (us). CORRIENTES: Corrientes, 4 Sept. 
1959, Mattson 3119 [grown in Copenhagen, collected by Pedersen in 
1953] (c). entre Rios: Concepcion del Uruguay, 3 Dec. 1877, 
Lorentz (gh). Uruguay. RIVERA: Cunapiru, 6-700 ft., 1928, Wing Jit 
(bm). CERRO largo: Rio "Rranco, 5-10 m., 27-29 Nov. 1947, H crier 
2127 (mo, NY, us), treinta Y tres : Vergara, 20 m., Dec. 1932, 
Hcrter 1608 (us). 

Tripogandra glandulosa has very distinctive seeds and 
this fact was recognized by the authors of the names 
Tradescantia glandulosa, T. radiata, DescantaHa pflanzii. 
While the seeds are diagnostic, other statements in all of 
the original descriptions are somewhat misleading. An 
examination of the type specimens shows all the floral and 

seed characters to be identical. 

The combination Tripogandra pflanzii has been published 
three times. The Gray Herbarium Card Index credits 
Celarier (1955) with the earliest combination. This com- 
bination was not validly published according to Article 33 
of the International Code of Botanical Nomenclature 
(1966). Burkart also combined the epithet in 1959, but 
this publication was later than Rohweder's (1956) valid 
publication of the name. 

Bacigalupo (1964) was the first person to consider 
Tripogandra radiata and T. pflanzii the same species but 
he does not seem to have seen the type specimen of T. 
glandulosa and recognized its identity. 

Several collections of apparently sterile plants have 
been made in Trinidad {Broadway 9145 (BM, uc) ; Fend- 
Jer 862 (bm, k) ; Grisebach 22 (k)], French Guiana 
[Sag. 948 (bm)], and Para, Brazil [Ruber 93 (bm)]. 
Pollen from an herbarium specimen mounted in aniline 
blue-lactophenol did not stain and is interpreted as having 
been sterile and non-functional when fresh. No specimens 
have seeds though inflorescences are present. 

These plants differ in the generally larger leaves (to 7.5 
cm long), the presence of one to several lines of hairs on 



268 



Rhodora [ Vo1 - 77 



mid 



-^ w 

leaf surface, peduncles to 4.4 cm long-, and all staminal 
filaments glabrous. Broadway's collections have small 
stolon-like branches perforating the leaf sheaths. 

The plants have been included in Tripogandra glandulosa 
because the morphological similarities are greater than 
with any other species. This decision is made reluctantly 
because it violates the morphological and geographic unity 
of the main body of collections. These plants seem to 
represent disjunct outliers of the main group of popula- 
tions of T. glandidosa and with the apparent sterility, 
warrant further collection and study. 

9. Tripogandra grandiflora (Donnell-Smith) Woodson, 

Ann. Missouri Bot. Gard. 29:153. 1942. 
Callisia grandiflora Donnell-Smith, Bot. Gaz. 31 :125. 
1901. type: Guatemala, alta verapaz: Cubilguitz, 
350 m., Apr. 1901, von Tuerckheim 7684 (Lectotype, 

us!: isolectotypes, gh!, mo!, us!). 

DonneUia grandiflora (Donnell-Smith) Clarke in Don- 
nell-Smith, Bot. Gaz. 33:261. 1902. 

Neodonnellia grandiflora (Donnell-Smith) Rose, Proc. 
Biol. Soc. Wash. 19:96. 1906. 

Illustrations: Donnell-Smith, Bot. Gaz. 33: PI. XL 
1902: Moore, Jr., Baileya 8:78-80, Figs. 24, 26. 1960. 

Plant perennial; stem first erect and later trailing over 
shrubs, to 3 m tall, to 1.0 cm in diam., unbranched or 
branched: internodes to 17 cm long, green, glabrous or 
with a complete or partial line of uniseriate hairs extend- 
ing down the internode from the sheath above. Leaves 
two-ranked, narrowly ovate or elliptic, smaller apically and 
reduced to sheaths in the inflorescence; blades to 15.2 cm 
long, 4.4 cm wide, oblique at the base, cuneate on one side 
and rounded on the other, glabrous dorsally, ventral sur- 
face glabrous except for the midrib which is wholly or 
partially covered with uniseriate hairs, margin ciliate or 
entire with a few hairs near the sheath, apex acute ; sheaths 



1975] Tripogandra — Handlos 269 



of vegetative leaves to 16 mm long, 8,0 mm in diam. when 
dry, villous at the orifice, otherwise glabrous except for a 
complete or partial line of uniserate hairs extending down 
the side opposite the blade, sheaths of the cataphylls to 
28 mm long, to 13 mm in diam., margin entire or ciliate. 
Inflorescences of a single terminal double cincinnus or 
paniculate, sometimes flexuous, composed of as many as 
9 terminal and axillary double cincinni ; peduncles to 4.0 
cm long, or so reduced that cincinni appear sessile, green 
or dark green with lighter green flecks, glabrous or with 
1 or 2 lines of uniseriate hairs; double cincinni with up 
to 13 buds, flowers and/or fruits; pedicels to 14 mm long 
at anthesis, to 1.3 mm in diam., green or green with a 
reddish tinge, glabrous, erect in fruit; bracts at the base 
of each pedicel entire, glabrous or with a few marginal 
uniseriate hairs. Flowers w 7 hite; sepals narrowly ovate or 
elliptic, to 8.5 mm long, to 4.0 mm wide at anthesis, green 
or green with a pinkish base, glabrous, margin entire and 
hyaline, apex acute or obtuse; petals ovate, elliptic, or 
obovate, to 10.5 mm long, to 8.1 mm wide, cuneate at the 
base, apex obtuse ; stamens 3, opposite the sepals, filaments 
1.0-2.5 mm long, white, glabrous (Fig. 9), anthers 0.9-1.3 
mm long, 0.8-1.2 mm wide, whitish, dorsifixed, versatile, 
open anther sacs white with a purple line around the edge 
(Fig. 8), pollen white; staminodes 3, opposite the petals, 
slightly epipetalous, filaments to 8.0 mm long, white, with 
two patches of uniseriate hairs in the upper half, the 
higher patch on the ventral side, the lower patch on the 
dorsal side with a few scattered hairs below the patch, 
filament bent in an open S-shape (Fig. 28), anthers dorsi- 
fixed, not versatile, connective orange or yellow, C-shaped 
with the anther sacs divergent, or connective elongate 
with anther sacs parallel, pollen yellow, sterile; ovary 0.7- 
1.6 mm long, 0.8-1.3 mm in diam., white or with a line of 
pink between the carpels, glabrous, style to 1.1 mm long, 
stigma simple or capitate and 3-lobed. Capsule elliptic, 
green or brown, glabrous, to 6.5 mm long; seeds gray or 
white and roughened, 1 or 2 per locule, elliptic (Fig. 77) 



270 



Rhodora [Vol. 77 



or triangular (Fig. 76), when elliptic with two sides up- 
turned above the protuberant embryotega, to 5.0 mm long, 
when triangular with two sides and one end upturned 
forming an angular C-shaped ridge around the embryotega, 
to 3.0 mm long, hilum linear (Fig. 78). 

Chromosome number: 7i=8, 16. 

Vernacular name: hoja de fluxion, Standley and Steyer- 
mark, Guatemala, Fieldiana: Bot. 24(3) :38. 1952. 

Distribution and habitat : the Yucatan peninsula, south- 
ern Mexico, Guatemala and British Honduras; at low 
elevations to 800-1500 m (7000-8800 ft, Nelson 3236c). 



Flowering: 



September 



in the area surrounding Tuxtla Gutierrez in Mexico and 
coincides with the rainy season in this location. In the 
area of Yucatan flowering occurs later, January to March 
and April. The fragrant flowers open sometime before 
sunrise and remain open until noon. 

REPRESENTATIVE SPECIMENS 

Mexico. Chiapas: km. 1026 of Hwy. 190, 24 km. west of Ocozo- 
cuautla, 620 m., 13 Au#. 1967, Handles 342 (bii). CAMPBCHE: Mon- 
terrey, 22 Jan. 1932, Lundell 1225 (ds, f, MICH, us). British Hon- 
duras. EL CAYO: Augustine, Mountain Pine Ridge, 1450 ft., 18 Mar. 
1960, Hunt 399 (bm, MICH, MO, us). Guatemala, peten: Santa Cruz, 
27-28 Mar. 1931, Bartlett 12390 (f, mich, us), alta verapaz: be- 
tween Coban and Finca Chimote, near Rubeltein, 800-1500 (300- 
500) m., 25 Feb. 1942, Steyermark 44203 (F, mo), izabal: Rio 
Chacon, 300 ft, 10 Feb. 1921, Johnson 1266 (us). 

Two different morphological forms can be distinguished 
within Tripogandra grandiflora. One form, representing 
the typical element, is more wide-spread and is charac- 
terized by leaves with a ciliate margin, a paniculate 
inflorescence, often with a flexuous axis and bearing 3-9 
double cincinni, staminodes with a C-shaped connective and 
divergent anther sacs and commonly one seed per locule. 
Based on information on specimen labels the flowers are 
very fragrant. This form is found in the eastern portions 
of Guatemala, British Honduras, and southeastern Mexico, 
the Yucatan peninsula and Chiapas. 



1975] Tripogandra — Handlos 271 



The other form has leaves which lack the ciliate margin, 
there being only a few hairs at the base of the blade near 
the sheath. The inflorescence is composed of one to three 
double cincinni, the staminodes have an elongate, straight 
connective and the anther sacs are parallel. There are one 
or two seeds per locule. The flowers have a weaker fra- 
grance if Handlos 342 is typical. While the evidence is 
limited there may also be a difference in a staminodial con- 
nective color, this form being yellow while the former is 
yellow or orange. The sterile pollen is more elongate in 
the typical element and more similar to the fertile pollen 
in Handlos 342. The second form is found in a restricted 
area in Chiapas, Mexico, centering around Tuxtla Gutierrez 
(Ocozocuautla to San Cristobal de las Casas). 

At the present time the two forms seem to be undergoing 
independent evolution. There may be significant inter- 
actions and relationships between the pollinators and such 
flora] characters as form and color of staminodes, and 
fragrance. If the two forms evolve in such a fashion that 
two distinct sets of pollinators are involved in pollination 
these two entities could retain their identity in the future 
if the populations become sympatric. Geographical isola- 
tion with some slight morphological and ecological differ- 
entiation now seems to exist. With further increase in 
differences these forms could be recognized as species but 
at this point in time it seems premature to give any nomen- 
clatural recognition to these forms. 

The United States National Herbarium is the location 
of two specimens which are held as the type material. 
According to the International Code of Botanical Nomen- 
clature, Article 7, Note 3, only one specimen can serve 
as the holotype of the species. I designate the sheet (us 
936917) which was annotated by C. V. Morton in August, 
1940, as Neodonnellia grandiflora, as the lectotype with 
the other sheet (us 936916) to serve as an isolectotype. 
I have chosen this particular sheet because it holds both 
flowering and fruiting branches. 



272 Rhodora [Vol. 77 



This species is distinguished by the revolute seed mar- 
gin, the farinose seed surface, the two patches of hairs on 
the inner filaments, the elongate shape of the cells of the 
staminodial hairs and the sweetly scented flowers. 

Reproduction in Tripogandra grandiflora is probably of 
two sorts, sexually by seeds and vegetatively from sections 
of fallen stem. Old stems fracture easily and in Mexico 
T have found young plants attached to pieces of stems 
lying on the ground; the axillary bud at the base of the 
internode apparently began to grow when conditions be- 
came favorable, i.e., when sufficient moisture was available. 



mav 



stem 



to a new location. In areas with abundant tropical showers, 
pieces of stem could be carried and spread to new loca- 
tions by water rushing down ravines and gullies. 

10. Tripogandra guerrerensis Matuda, Anales Inst. Biol. 

Nac. Mexico 36:113. 1966 ('1965'). TYPE: Mexico. 
Guerrero: Rincon de la Via, 775 m., 24 Sept. 1961, 
Krvse 461 (Holotype, mexu; isotype, mexu!). 

Illustration: Matuda, Anales Inst. Biol. Nac. Mexico 
36:113. 1966 ('1965'). 

Plants annual; stems erect, to 74 cm tall, 2-4 mm in 
diam. when dry, unbranched or with 1-2 branches basally ; 
internodes 1.8-9.7 cm long, green, glabrous except for a 
line of uniseriate hairs continuing down the internode 



from 



Leaves nar- 



rowlv ovate; blades 2.9-10.7 cm long, 0.3-1.1 cm wide, 
narrowed toward the base, smaller distally and leaves 
reduced to sheathing bracts in the inflorescence, glabrous 
ventrally, pilose dorsally, margin ciliate, apex acute; 
sheaths 2.0-12 mm long, villous at the orifice, otherwise 
glabrous except for a line of uniseriate hairs which con- 
tinues down the side of the sheath opposite the blade. 
Inflorescences paniculate, much-branched, terminal and in 
the axils of the upper leaves, composed of 7-30 double cin- 



1975] Tripogandra — Handlos 273 



cinni, peduncles 7-23 mm long*, 0.4-0.6 mm in diam., green, 
glabrous; bracts within the inflorescence with short lamina 
or reduced to a sheath near the apex ; buds, flowers and/or 



mm 



mm in diam 



uniseriate or capitate hairs, erect in fruit; bracts sub- 
tending the pedicels glabrous, margin entire. Flowers 
white; sepals ovate-cymbiform, 2.3-3.2 mm long, 1.5-2.1 mm 
wide, green with purple tips, glabrous or with a few scat- 

■ 

tered capitate hairs, margin entire, apex acute or obtuse; 
petals ovate-elliptic, 3.5-5.0 mm long, 2.3-3.5 mm wide., 
tapering at the base, apex obtuse; stamens 3, opposite the 
sepals, filaments 1.0-1.1 mm long, white, glabrous basally 



m 



mm 



colorless hairs (Fig. 11), anthers 0.6-0.7 
mm wide, white, basifixed, versatile with anther sacs 
parallel, longer than the gibbous connective, pollen white ; 
staminodes 3, opposite the petals, filaments slightly epipe- 



mm 



in the upper half, expanded in the bent portion (Figs. 15, 
17), anthers dorsifixed, versatile, filaments attached near 
one side of the yellow, discoid connective and anther sacs 
borne on the side opposite the filament, connective 0.6-0.8 
mm long, 0.5-0.7 mm wide, pollen yellow, sterile ; ovary 
0.6-0.8 mm long, 0.6-0.8 mm in diam., light green, glabrous, 
style 0.2-0.3 mm long, stigma simple, papillate. Capsule 
obovoid, glabrous, light brown, 2.9-3.6 mm long, 2.2-2.5 
mm in diam.; seeds 2 per locule (Fig. 49), the upper in 
each locule longer (Figs. 70, 71), 1.6-1.9 mm, the lower 
shorter, 1.1-1.4 mm (Figs. 68, 69), triangular to elliptical, 
dark brown to black, testa areolate with ribs radiating 
from the embryotega, hilum linear-elliptical (Figs. 69, 71). 
Chromosome number: ?? = 8. 

Distribution and habitat: Mexico in the states of Jalisco 
and Guerrero; in rocky areas at low elevations. This species 
is known from two locations which are about 455 km apart. 
The intervening area is not easily accessible but it seems 
highly probable that there are other suitable habitats be- 



274 Rhodora [Vol. 77 



tween the two known locations, and as the area is explored 
new collections will almost certainly be made. 

Flowering: Flowering occurs in September and October 
in the two locations where this species has been collected. 



Mexico 



AM and close about 2:30 PM 



SPECIMENS EXAMINED 

Mexico. JALISCO: just before km. 148 on Hwy. 110 to Colima, 
5.6 kin. south of bridge over Rio San Pedro, ca. 1050 m., 2 Oct. 1967, 
Handlos 438 (bh); km. 147-148 on road between Colima and Ma- 
zamitla, 1040 m., 10 Sept. 1961, Moore, Jr. & Bunting 8746 (bh, uc). 

Tripogandra guerrerensis is unique among the species 
I have seen in Mexico because the petals are not sym- 
metrically disposed at anthesis; rather, two petals bend 
upward so that an angle of approximately 90° is formed 
between the middle upright petal and the two lateral ones, 
whereas in all the other species the angle between petals 
is about 120°. 

The androecium bears a constant relationship to the 
petals. In typical Tripogandra-f&shion, the filaments of 
two of the staminodes bend around two staminal filaments 
and the three staminodes are then positioned in front of 
the middle upright petal. The stamens surround the ovary 
but because of the position of the petals and staminodes 
the anthers can not be approached by a pollinator alighting 
on the petals. From observations made of a natural popu- 
lation in Jalisco, Mexico, on October 2 and 3, 1967, the 
disposition of the petals seems significant because bees are 
restricted in their movements and are able to gather pollen 
only by clinging to the staminodes. If the pollinator col- 
lects sterile pollen, it hangs from the filaments of the 
sterile stamens, with its abdomen pointed toward the 
center of the flower. While gathering sterile pollen, the 
abdomen is placed on the fertile anthers so that pollen is 
deposited on the pollinator's abdomen. This seems to be 
a device for insuring pollination. Insects are also able 
to collect fertile pollen by clinging upside down to the 



1975] Tripogandra — Handlos 275 



staminodes with the head near the anthers. Further 
critical studies are needed to determine how much cross- 
pollination and how much self-pollination is effected. In 
the greenhouse, abundant viable seed was set after insects 
ceased entering from outdoors, so the plants are not self- 
sterile. 

11. Tripogandra kruseana Matuda, Anales Inst. Biol. Nac. 

Mexico 37:77, 78. 1967 (1966'). holotype: Mexico. 
GUERRERO: Rincon de la Via, 735 m., 16 Oct. 1965, 

Kruse 903 (mexu!). 

Illustration: E. Matuda, Anales Inst. Biol. Nac. Mexico 
37:77. 1967 ('1966'). 

Plants perennial?, rooting at the nodes; flowering stems 
erect, to 61 cm tall, branched at the base; internodes to 
9.5 cm long, with a partial line of uniseriate hairs extend- 
ing down the side from the sheath above, otherwise 
glabrous in the lower portion of the plant. Leaves nar- 
rowly ovate: blades to 10.1 cm long, to 1.2 cm wide, 
glabrous, margin ciliate, apex long acuminate; sheaths to 
12 mm long, to 1.0 cm in diam., a few long hairs at the 
orifice, with a line of uniseriate hairs on the side opposite 
the blade, otherwise glabrous. Inflorescences borne termin- 
ally and in the upper 3 leaf axils, composed of up to 9 
double cincinni; peduncles to 1.3 cm long, pilose with 
capitate hairs; buds, flower and/or fruits 2-3 per double 
cincinnus; pedicels 8-12 mm long, green, erect in fruit, 
pilose with capitate hairs; bracts at the base of each 
pedicel with margin entire or denticulate. Flowers pink 
(white fide label) ; sepals ovate, 3.1 mm long, 1.6 mm wide 
in flowers available (2 mm long, 1.2 mm wide fide desc), 
pilose with capitate hairs, margin red; petals not avail- 
able; stamens 6, in two whorls, 3 shorter, 0.7 mm long, 
bearded with white moniliform hairs; 3 larger, 2 mm 
long, bearded with white moniliform hairs; ovary green, 
glabrous, style short, stigma capitellate. Capsule globose, 
2.5 mm long, 2.5 mm in diam. (immature). 



276 



Rhodora [Vol. 77 



Distribution and habitat: This species is known only 
from the type collection at Rincon de la Via, Guerrero, 
Mexico, in a rocky area at 735 m. altitude. 



Flowering: The type specimen was in flower in mid- 
October. 

Matuda's original description of T. kruseana describes 
the base of the leaf as amplexicaul but this is not readily 
discernible in the holotype. If the plant has amplexicaul 

* 

leaves and is a perennial, it will key out to T. e?icolea. If 



» 



the plant lacks amplexicaul leaves, as the photograph of 
the holotype seems to indicate, it will key near T. saxicola 
from which it may be distinguished by erect pedicels and 
pilose peduncles. 

■ 

12. Tripogandra montana Handlos, sp. hot. holotype: 

Mexico. CHIAPAS: 3.2 miles N of junction of Hwy. 
195 with road to El Bosque and Simojovel, 1770 
(1690) m., 15 Aug. 1967, Handlos 355 (us!). 

Herba perennis; caulis decumbens, usque ad 2.6 m 
longus, internodiis usque ad 12.8 cm longis, glabris vel 
raro distaliter pilosis, linea uni'ca pilorum instructis. 
Folia angusto-ovata, laminis usque ad 14.5 cm longis, 
usque ad 3.5 cm latis, apice acutis, basi obliquis, glabris 
vel pilis paucis dispersis instructis vel pilosis, margine 
ciliatis vel medio nonnunquam eciliatis, vaginis usque ad 
18.0 mm longis, usque ad 12.5 mm diam., linea unica 
pilorum instructis, aliter glabris vel pilis dispersis in- 
structis vel pilosis, orificio villosis vel pilis dispersis prae- 
ditis. Inflorescentiae terminales et in 1-4 axillis foliorum 
summorum, ex 1-17 cincinnis duplicibus constantes; 
pedunculi 0.7-12.7 cm longi, glabri, pilosi vel 1-2 lineis 
pilorum instructi ; cincinwi duplices omnes alabastra, flores, 
vel fructus usque ad 17 gerentes; pedicelli usque ad 2.5-6.0 
mm longi, 0.6-0.8 mm diam., maturitate reflexi, pilis capi- 
tals paucis instructi vel pilosi, bracteis basi pedicellorum 
glabris vel pilis capitatis vel uniseriatis dispersis pubes- 
centibus, margine ciliatis vel eciliatis. Flores rosei ; sc))ala- 



1^5] , Tripogandra — Handlos 277 



anguste ovata, elliptica, vel obovata, cymbiformi 



mm 



capitatis instructa, margine integra et hyalina, apice 



mm 



basi cuneata, apice rotundata; stamina 6 in verticillis 
duobus, 3 sepalis opposita filamentibus brevibus, 1.0-2.5 
i^im longis, subroseis, glabris vel raro pilis paucis praeditis 
(Fig. 10), antheris 0.8-1.8 mm longis, 0.6-1.2 mm latis, 
polline albido, 3 petalis opposita filamentibus epipetalis, 
sigmoideis, 4.5-7.5 mm longis, medio pilis roseis monili- 
formibus pubescentibus (Fig. 31), antheris 0.5-0.9 mm 



mm 



glabrum 



stigmate capitellato vel capitato. Capsula elliptica, 2.0-3.7 
mm longa, 1.6-2.6 mm diam., glabra, seminibus triangulari- 
bus, 1.1-1.7 mm longis, reticulatis (Figs. 44-46), hilo 
punctiformi (Fig. 45). 

Chromosome number: n~21. 

Vernacular name: tzima fide Standley, Guatemala. 

Distribution and habitat: southern Mexico, Guatemala, 
Honduras, and El Salvador; in pine-oak forest areas at 
higher elevations, ca. 1100-2500 m. The one undisturbed 
location in which I found this species was in a black loam 
soil in a pine woods with a scattering of oaks, Liquidambar 
and tree ferns. The plants grew on a moist, east-facing 
slope. An adjacent, drier, sunnier and steeper west-facing 
slope had no plants of this species. 

Flowering: In Mexico flowering occurs between March 
and November; in Guatemala between November and June; 
in Honduras in February, March, April, July and Novem- 
ber. The flowers open about 8:00 AM in their natural 
habitat and close about 2 :00 PM. The plants I found had 
odorless flowers. Flowering seems to extend over a long 
period of time because both young inflorescences and 
mature seed were found in the location where Handlos 
355 was collected. 



278 



Rhodora t Vo1 - 77 



4 






REPRESENTATIVE SPECIMENS 

Mexico. Chiapas: in the paraje of Kulak'tik, Municipio of Tene- 
japa, 5500 ft, 25 Nov. 1965, Breedlove 14205 (ds, F, mich). Guate- 
mala. HUEHUETENANGO : Canyon of Rio Chixoy near Malacatancito 
about 20 km. southwest of Huehuetenango, 1600 m., 1 Dec. 1962, 
Williams, Molina R. cfe Williams 22149 (F, NY). QUICHE: Cunen, 6000 
ft, Apr. 1892. Heydc & Lux 3521 (gh, m, ny, us), alta verapaz: 
Chicoyonito, 4300 ft., Apr. 1889, Donnell-Smith 1643 (gh, ny, ph, 
us), san MARCOS: slopes bordering Rio Malacate, barrancos 6 miles 
south and west of town of Tajumulco, northwestern slopes of Volcan 
Tajumulco, 2300-2800 ,m., 26 Feb. 1940, Steyei~mark 36679 (f). 
QUEZALTENANGO: mountains above Rio Samala, Sierra Madre Moun- 
tains, 2 km. west of Zunil, 2300 m., 14 Dec. 1962, Williams, Molina R. 
& Williams 23023 (F, ny). solola: steep slopes of Panajachel 
water falls, road to Solala, 2200 m., 12-23 Jan. 1966, Molina R., 
Burger & Wallcnta 16233 (F, ny). chimaltenango: near Rio 
Pixcayo, between Chimaltenango and San Martin Jilotepeque, 
1650-1800 m., 3 Feb. 1939, Standley 64362 (f). sacatepequez : 
slopes of Volcan de Agua, south of Santa Maria de Jesus, 1800-2100 
m., 10 Dec. 1938, Sta'ndley 59449 (f). Guatemala: damp wooded 
barranca 10 km. south of San Raimundo, about 1800 m., 18 Jan. 1939, 
Standley 62882 (f). EL progreso: Montana Canahui, between Finca 
San Miguel and summit of mountain, near upper limits of Finca 
Caieta, 1600-2300 m., 10 Feb. 1942, Steyermark 43764 (f). zacapa: 
along Rillito del Volcan de Monos, Volcan de Monos, 1150-2100 m., 
10 Jan. 1942, Steyermark 42336 (f, mo), chiquimula: Montana 
Nono.ia, 3-5 miles east of Camotan, 600-1800 m., 11 Nov. 1939, 
Steyermark 31687 (f). suchitepequez: southwestern lower slopes 
of Volcan Zunil, in vicinity of Finca Montecristo, southeast of Santa 
Maria de Jesus, 1200-1300 m., 31 Jan. 1940, Steyermark 35221 (f). 
Honduras, copan: 5 km. al S.O. de Santa Rosa de Copan, 1200 m., 
29 Mar. 1963, Molina R. 11675 (f). comayagua: vicinity of Sigua- 
tepeque, 1080-1400 m., 14-27 Feb. 1928, Standley 56198 (F, us). 
intibuca: Bahos de Esperanza, 1800 m., 27 Nov. 1958, Hawkes, 
Hjerting & Lester 2103 (c, F). El .Salvador. Santo Tomas, 1922, 
Calderon 1294 (ny, us). 

The collections which I have included in Tripogandra 
montana have usually been identified as T. elongata in the 
past. On the basis of a distinct seed reticulation, a capitate 
stigma, a short style, a different leaf texture, adaptation 
to a relativelv moist montane habitat, and a different and 
distinct geographic range, I must recognize T. montana 
as different from T. elongata. Tripogandra elongata is 



1975] Tripogandra — Handlos 279 



discussed further under T. serrulata. The long peduncles 
and bright pink flowers of T. elongata may indicate some 
affinity with T. montana but this is only speculation. 

Variation in leaf vesture does occur within this taxon 
but that seems to be a common occurrence within this 
genus. The significance, function, and mode of inheritance 
of this character should be investigated further. 

13. Tripogandra multiflora (Swartz) Rafinesque, Flora 

Telluriana 2:16. 1837 ('1836'). 

Tradescantia multiflora Swartz, Nova genera & species 
plantarum seu prodromus . . . Indiam occidentalum 
57. 1788; Flora Indiae occidentals 1:599. 1797; 
3:1972. 1806; Jacquin, Collectanea ad botanicam 
. . . 3:226, 227. 1791. type: Jamaica. Swartz 



(Holotype, (b)?; isotype, m!). 

Tradescantia, parviflora Ruiz & Pavon, Florae Peru- 
vianae et Chilensis prodromus, . . . 3:43. 1794. 
type: Peru, huanuco: Pozuzo, 1778-1788, Ruiz 
& Pavon. (Holotype, ma; photograph, ma!; iso- 
type, f!). 

Tradescantia procumbens Willdenow, Species Planta- 
rum 2:19 1799. HOLOTYPE: Jacquin (b) ; photo- 
graph, b!. 

Commelina floribunda HBK., Nova genera et species 
plantarum 1:260. 1816. holotype: Cumana, Bor- 
dones & Quetepe, Humboldt & Bonpland (b) ; photo- 
graph, b!. 

Heminema multiflora (Swartz) Rafinesque, Flora 
Telluriana 2:17. 1837 ('1836'). 

Aneilema floribunda (HBK) Hooker & Arnott, The 
Botany of Captain Beechey's Voyage 311. 1840. 

Tradescantia cumanensis Kunth, Enumeratio Planta- 
rum 4:96. 1843, based on Commelina flonbunda 
HBK non Tradescantia flonbunda M. & G. (1842) 
nee T. floribunda Kunth (1843). 

Tradescantia multiflora y linnaei Clarke in DC, Mono- 
graphiae Phanerogamarum 3:306. 1881. type: 






280 



Rhodora [Vol. 77 



Colombia, in montibus juxta Bogotam, 6 Nov. 
1852, Holton 127 (Holotype, k!; isotype, ny!). 

Trade scant ia multiflora ft parvi flora (Ruiz & Pa von) 
Clarke in DC, Monographiae Phanerogamarum 

3:306. 1881. 

Descantaria procumbens Schlechtendal fide Hasskarl 
ex Clarke in DC, Monographiae Phanerogamarum 

3:305. 1881, nom. nud. pro syn. 

Leptorhoeo floribunda (HBK) Baillon, Histoire des 
Plantes 13:218. 1894 ('1895'). 

Tradescantia ionantha Diels in Urban, Bot. Jahrb. 
Syst. 37:382. 1906. type: Peru, puno: Sandia, 
2100 m., 24 Mar. 1902, Weberbauer 588 (Holotype, 
b!; isotype, g!). 

Tradescantia multiflora var. tobagensis Urban, Sym- 
bolae Antillanae 7:174. 1912. holotype: Tobago. 
juxta flumen "Bacolet" ad "Calder Hall", 31 Oct. 
1889, Eggers 5693 (b!). 

Descantaria multiflora (Swartz) Bruckner, Notizbl. 
Bot. Gart. Berlin-Dahlem 10:56. 1927. 

Descantaria cumanensis (Kunth) Schlechtendal ex 
Bruckner, Notizbl. Bot. Gart. Berlin-Dahlem 10:56. 
1927. 

Descantaria ionantha (Diels) Bruckner, Notizbl. Bot. 
Gart. Berlin-Dahlem 10:56. 1927. 

- 

Tripogandra floribunda (HBK) Woodson, Ann. Mis- 
souri Bot. Gard. 29:152. 1942. 

Tripogandra cumanensis (Kunth) Woodson, Ann. Mis- 
souri Bot. Gard. 29:152. 1942. 

Tripogandra ionantha (Diels) Macbride, Revista Univ. 
(Cuzco) 33(87) :142. 1945. 

Tripogandra parvi flora (Ruiz & Pa von) Steyermark, 
Phytologia 9:339. 1963 ('1964'). 

Tripogandra midtiflora (Swartz) Woodson [fide Baci- 
galupo] forma parvi flora (Ruiz & Pavon) Baciga- 
lupo, Darwiniana 14:398. 1967. 



1075] Tripogandra — Handlos 281 



Illustrations: Jacquin, Tcones Plantarum rariorum 2: 
355. 1790 ('1781-1795') ; Moore, Jr., Baileya 8:81, Fig. 27. 
1960: Bacigalupo, Darwiniana 14:399, Fig. 1, 1967. 



Plants perennial, trailing and rooting at the nodes or 
caespitose; floriferous stems upright, to 83 cm tall, un- 
branched or branched; internodes to 13.5 cm long, the 
uppermost 1-4 pilose (rarely glabrous), the lower glabrous 
or glabrous proximally and pilose distally, in all speci- 
mens with a line of uniseriate hairs extending down the 
stem from the sheath above. Leaves narrowly ovate, ovate 
or broadly ovate; blades to 8.8 cm long, to 2.8 cm wide, 
glabrous or pilose dorsally, the ventral surface variously 
glabrous, pilose and/or w T ith a line or band of uniseriate 
hairs near the margin, occasionally the base of the midvein 
with a line of uniseriate hairs, base oblique, cuneate, or 
rounded, margin ciliate, apex acute; sheaths to 14.2 mm 
long, to 9.2 mm in diam., villous at the orifice, the upper- 
most 1-7 pilose (rarely glabrous), the lower ones glabrous 
or pilose, in all specimens with a line of uniseriate hairs 
extending down the side opposite the blade. Inflorescences 
of 1-26 double cincinni borne terminally and in the axils 

t 

of the upper 1-8 leaves; peduncles to 3.5 (-5.4) cm long, 
densely pilose or rarely pilose distally and glabrous prox- 
imally; double cincinni with up to 16 buds, flowers and/or 
fruits; pedicels 1.4-4.0 mm long, reflexed in fruit, glabrous, 
with scattered uniseriate hairs or pilose; bracts at the 
base of each pedicel glabrous, with scattered uniseriate 
hairs, or pilose, margin entire or long-ciliate. Flowers 
white or pink; sepals ovate-cymbiform, 1.8-4.0 mm long, 
1.0-1.8 mm wide, green, red, or green with a red tip and/or 
base, glabrous, with scattered capitate hairs, or pilose, 
margin hyaline, apex more or less acute ; petals broadly 
ovate (?), 1.6-4.0 mm long, 1.0-ca. 2.2 mm wide; stamens 6, 
in two whorls, the outer shorter, filaments 0.5-1.3 mm long, 
glabrous, anthers 0.3-0.6 mm long, 0.3-0.6 mm wide, pink, 
basifixed, anther sacs parallel; stamens of the inner whorl 
longer, 1.4-3.2 mm long, S-shaped, bearded with a ventral 



282 



Rhodora [Vol. 77 



tuft of moniliform hairs below the expanded distal end, 
anthers 0.3-0.5 mm long, 0.2-0.8 mm wide, dorsifixed, con- 
nective I- or C-shaped, yellow, anther sacs divergent or 
parallel through bending of the connective; ovary 0.5-1.0 
mm long, 0.4-0.8 mm in diam., glabrous, style 0.15-0.3 mm 
long, stigma capitellate or capitate. Capsules 1.4-2.5 mm 
long, 1.5-2.5 mm in diam., green or light brown, glabrous; 
seeds 1-2 per locule, rounded-triangular, 0.7-1.0 (-1.4) mm 
long, light or dark gray or brown, testa reticulate-foveate 
(Figs. 32, 34), hilum punctiform (Figs. 33, 35). 

Chromosome number: 2w— 64 (A. Sparrow, pers. 
comm.) 

Distribution and habitat: Jamaica, Trinidad, Tobago, 
Costa Rica, Venezuela, Colombia, Peru, Bolivia, and Argen- 
tina; on shaded or open rocky banks or slopes from sea 
level to 2900 m. 

Flowering: In Jamaica flowering occurs from June to 
February; in Trinidad and Tobago in July and October; 
in Costa Rica in November; in Venezuela from August to 
January and in May and June; in Colombia from May to 
February; in Peru in March, July, November, and Decem- 
ber; in Bolivia from December to May. 



REPRKSENTATIVE SPECIMENS 

Jamaica. Troy, 2000 ft., 1G Oct. 1917, Harris 12573 (F, NY, us). 
Manchester: vicinity of Mandeville, 15-26 Feb. 1910, Brawn 84 
(NY, ph). ST. anurew: off road from Kingston to Newscastle, 1250 
ft., 17 June 1963, Crosby, Hespevheide & Anderson 178 (F, MICH, 
msc, uc). Tobago. Roxboroujyh, 20 July 1914, Broadway 4906 (us); 
Mason Hall, 19 Oct. 1937, Sandwith 1865 (ny). Trinidad. Point 
opposite Melville Island, 25 Oct. 1925, F[reeman] & W[iUiams] 11408 
(ny). Costa Rica. SAN JOSE: vicinity of El General, 1160 m., Nov. 
1936, S hutch 2935 (gh, k, mo, ny, us). Venezuela, falcon: Mea- 
cbiche, 18 Jan. 1942, Lasser 178 (us, yen), lara : Loma de Leon 
Dto. Yribarren, 16 Sept. 1950, Taniayo 3726 (ven). distrito federal: 
La Quesera, Upper Cotiza, near Caracas, above 1400 m., 18 Sept. 
1921, Pitticr 9821 (us, ven). Miranda: Los Blanches, on new road 
to Sta. Lucia, 14 Pec. 1924, PHtier 11636 (us, ven). merida: 3 
miles west; 2 miles south of Merida Valley, 4500 ft., 26 Jan. 1931, 
Reed 335 (us). Colombia, magdalena: Sierra Nevada de Santa 
Marta, southeastern slopes; Hoya del Rio Donachui; below the vil- 



J5>75] Tripogandra — Handlos 283 



la^e Donachui near the river, 1350-1230 m., 24 Sept. 1959, Cuatre- 
casas & Castaneda 24406 (us), norte de santander: vicinity of 
Mutiscua, 2900 m., 20-22 Feb. 1927, Killip & Smith 19669 (us). 
santander: vicinity of Charta, 2000-2600 m., 1-11 Feb. 1927, Killip 
& Smith 17446 (gh, ny, us), boyaca: Valle de la Uvita, Cordillera 
Oriental, 2490-2560 m., 16 Sept. 1938, Cuatrecasas 1857 (us). 
CUNDINAMARCA : Macizo de Bogota, Quebrada de Chico, 2650-2750 
m., 1 June 1939, Cuatrecasas 5224 (us). HUILA: Cordillera Oriental, 
east of Neiva, 800-1000 m., 31 July 1917, Rusby & Pertnell 424 (ny). 
Peru, san martin: prope Tarapoto, 1855-6, Spruce 4198 (bm, br, 
E, ny). huanuco: west and above Puente Durand, north of Huanuco, 
1900 m., 4 Nov. 1938, Stork & Horton 9589 (G, gh, k). lima: Lima, 
1 July 1914, Rose & Rose 18551 (us). CUSCO: Potrero, Convencion, 
1300 m., 2 Mar. 1940, Vargas C. 1832 (git), puno: Sandia, 2100 
m., 24 Mar. 1902, Weberbauer 588 (b, g). Bolivia, la PAZ: Apolo, 
4800 ft., 10 Mar. 1902, Williams 55 (bm, us). 

Tripogandra multiflora is a widespread species in South 
America and a complex pattern of variation seems to exist 
within the species as I conceive it. Collections below about 
2,000 meters from Colombia, Venezuela and the Caribbean 
islands form a fairly uniform group characterized by a 
trailing habit, glabrous leaves with an oblique base, white 
flowers and pilose sepals. 

Collections from Bolivia in contrast are quite variable. 
The leaf base is usually oblique and the base of the plant 
seems to be decumbent. The flower color may be either 
white or pink, sometimes even in the same collections, e.g., 
Buchtien 399. The leaf indument is variable — the leaves 
may be (1) glabrous dorsally with a band of hairs near 
the margin ventrally as in Williams 55; or (2) pilose on 
both sides as in Rusby 1369, Buchtien 2457, 4183 (f, gh), 
7190 (gh), Bang 603 (f, m, mich) ; or (3) pilose dorsally 
with a ventral marginal band of hairs as in Buchtien s.n., 
399 (US), 4183 (F), 7190 (NY), Bang 603 (E, P, GH, MO, 
ph, US). The sepals may be glabrous as in Williams 55, 
Buchtien s.n., 399 (us), 4183 (f), 5350, 7190 (ny), Bang 
603 (E, F, GH, MO, NY, PH, us), or have scattered capitate 
hairs as in Rusby 1369, Buchtien 399 (US), 2457 (ny), 
4183 (GH), 7190 (GH), Bang 603 (M, MICH) ; or be pilose 
as Buchtien 4183 (F, g). These plants must be studied in 



284 



Rhodora [Vol. 77 



the field for a better understanding of the problems. I do 
not know whether hybridization is occurring. It is possible 
that Elections have become mixed before being glued to 

* 

herbarium sheets; most collections were made by Buchtien 
and Bang. It is possible, however, that the species in 
Bolivia is very polymorphic with some obvious traits 
segregating* within the populations. 

Plants which grow in the highlands of Colombia, Peru 
and Bolivia, above 2,000 meters elevation, tend to be caes- 
pitose and the leaves are generally broadly ovate with a 
cuneate to nearly amplexicaul base. Compared to lowland 
plants of T. multiflora there are few inflorescences and 
these are partially covered by the upper one to five leaves ; 

- 

the sepals, petals and pistil tend to be longer and the 
stigma is broader, more nearly capitate than capitellate. 
The flowers are always bright pink or magenta. I can 
find no qualitative characters (aside, perhaps, from flower 
color) which will allow one to separate the plants of this 
group from the bulk of T. multiflora. The presence of 
plants which appear intermediate between the highland 



me 



as a mountain ecotype of T. multiflora. Intergradation as 
seen from herbarium specimens may be more apparent 
than real. It is possible that these forms are ecologically 
separated and generally do not intergrade but are very 
similar species. If one considers evolution to be a con- 
tinuous and gradual process and not saltational, then it 
is possible that the situation just described represents a 
slight ecological divergence with no overwhelming morpho- 
logical differences, a case of sibling or micro-species. These 
essential questions can only be settled with further field 
work. 

Because of the pattern of morphological variations oc- 
curring within Tripogandra multiflora, a number of names 
have been applied to this species. These names are con- 
sidered in detail here. 

Willdenow (1799) named Tradescantia procumbens from 
a collection with a procumbent stem from mainland South 



1!,T5 ] Tripogandra — Handlos 285 

America. This name seems unnecessary for there appear 
to be no other characters by which the island and the 
mainland populations differ. Observations of greenhouse 
grown plants and information on herbarium labels indi- 
cate that, in fact, the island plants also tend to trail over 
the ground. 

Ruiz and Pavon's Tradescantia parviflora does not seem 
sufficiently distinct to be recognized as a species, the only 
distinguishing character being the presence of pilosity on 
the dorsal leaf surface. Clarke (1881) considered T. parvi- 
flora a variety (/3) of Tradescantia multiflora. No other 
collections that I have seen duplicate the distribution of 
hairs found on the Ruiz and Pavon specimen. The other 
collections cited under variety parviflora by Clarke have 
additional hairs in a line near the margin on the ventral 
surface which are not found in the Ruiz and Pavon speci- 
men. Bacigalupo (1967) considered Tradescantia parvi- 
flora a form of Tripogandra multiflora. , The specimen 
which he cites from Argentina apparently has additional 
indument on the ventral leaf surface not found on the 
Ruiz and Pavon specimen. In terms of the distribution 
of hairs on the leaves, the Ruiz and Pavon specimen seems, 
therefore, to be an intermediate between the Bolivian, Ar- 
gentinian and other Peruvian collections and the Colombian, 
Venezuelan and Jamaican glabrous-leaved plants. Steyer- 
mark (1963) cited specimens of Tripogandra parviflora 
from Ecuador but apparently these are referable to a 
species of Gibasis. 

Urban's variety tobagevsis of Tradescantia multiflora 
is represented by a very few collections. While the seeds 
he observed were 0.6-0.7 mm in diameter, I find that seeds 
from other collections on Trinidad and Tobago range from 
0.7-1.0 mm in diameter. The plants do seem smaller but 
a formal Latin designation seems unnecessary. The small 
stature of the plants may represent only stunted growth 
and no real genetic difference. 

The mountain ecotype described earlier was named 
Tradescantia multiflora variety y linnaei by Clarke (1881) 



286 



Rhodora [Vol. 77 



and T. ionantha by Diels (in Urban, 1906). 



om 



rium specimens there seems 



no character ex- 



stem 



rare or at least 



cept, perhaps, the bright magenta flower color by which 
these plants can be distinguished. 

The names Tradescantia cumanensis and Tripogandra 
cumanensis have been applied to specimens of Tripogandra 
serrulata, but these usages seem to stem from an initial 
misidentification of specimens by Clarke. Kunth's original 
description of Tradescantia cumane7isis (as Commelina 
floribunda) notes that the top of the 
are "hirto-pilosis" while the sepals are "glanduloso-pilosis." 
These characters coincide with those found in T. multi- 
flora which is common in Venezuela where the type of 
T. cumanensis was collected, but these characters are not 
found in T. serrulata which seems 
not often collected in Venezuela. 

Kunth's concept of similarities may be considered as a 
further line of evidence to demonstrate the identity of 
T. multiflora and T. cumanensis. Kunth (1843) grouped 
similar species in his treatment of the family and he noted 
the affinities he believed existed. It can be seen that 
Tradescantia schlecht end 'alii (a synonym for Tripogandra 
serrulata) is number 41 in Kunth's list while T. procum- 
bens. T. parvi flora, T. multiflora, and T. cumanensis are 
numbers 44, 45, 46, and 47, respectively. Using this evi- 
dence as a reflection of similarities, T. cumanensis stands 
apart from T. schlechtendalii (= Tripogandra serrulata) 
but close to T. multiflora. I have only seen a photograph of 
the holotype of T. cumanensis but an examination of seeds 
of this specimen would make a positive indentification 

possible. 

By application of the Internal Code of Botanical Nomen- 
clature, Aneilema floribunda, Leptorhoeo floribunda and 
Tripogandra floribunda based on Commelina floribunda 
HBK must be cited in synonymy under Tripogandra multi- 
flora for reasons which follow. 

Hooker and Arnott published the combination Aneilema 
floribunda based on Commelina floribunda HBK but the 



1975 ] Tripogandra — Handlos 287 



specimen to which they applied this name was a species 
of Leptorhoeo, a genus in need of further study. Subse- 
quent authors, Baillon (1894) and Woodson (1942), have 
credited Hooker and Arnott as authors of the specific 
epithet, floribunda, but Hooker and Arnott correctly 
credited HBK and did not, in fact, publish a new species 
description though they did apply the name incorrectly. 
Rohweder (1956) published Tripogandra ionantha and 
Steyermark (1964) published T. multi flora as new com- 
binations. Both of these names were validly published 
earlier so neither Rohweder nor Steyermark can be credited 
as the correct authors. 

14. Tripogandra neglecta Handlos, sp. nov. holotype: 

Brazil, minas gerais: ad Lagoa Sta., 27 Mar. 1864, 
Warming 1069/1 (c!). 

Herba annua ( ?) ; caulis erectus, usque ad 40 cm altus, 
internodiis usque ad 10.5 cm longis, linea unica pilorum 
instructis aliter glabris. Folia angusto-ovata. 



ami 



cm loneris, usque ad 2.1 cm 



cuneatis, glabris, margine ciliatis, vaginis usque ad 2.3 cm 



cm 



* 7 

aliter glabris, orificio parce villosis. Inflorescentiae termi- 
nates et in axillis foliorum summorum, ex 4-9 cincinnis 
duplicibus constantes; pedunculi usque ad 4.0 cm longi, 
1-2 lineis pilorum instructi, cincinni duplices omnes ala- 
bastra, flores, vel fructus usque ad 12 gerentes; pedicelli 
usque ad 5.5 mm longi, glabri, maturitate reflexi, bracteis 
basi pedicellorum margine integris, glabris. Flores albi?; 
sepala ovata, cucullata, usque ad 4.0 mm longa, usque ad 
2.3 mm lata, pilis paucis uniseriatis instructa, margine 
integra et hyalina, apice ± obtusa; petala ovata, apice 
obtusa; stamina 6 in verticillis duobus, 3 sepalis opposita 
filamentibus brevibus, usque ad 1.3 mm longis, glabris, 
antheris usque ad 1.1 mm longis, usque ad 1.2 mm latis. 
3 petalis opposita filamentibus longioribus, usque ad 3.2 
mm longis, sigmoideis, barbatis distaliter, antheris usque 
ad 0.8 mm longis, usque ad 1.2 mm latis, connectivo C- 



288 



Rhodora [Vol. 77 



formi; ovarium usque ad 1.0 mm longum, usque ad 0.8 mm 
diam., glabrum, stylo usque ad 0.6 mm longo, stigmate 
simplici. Capsula globosa, usque ad 3.0 mm longa, usque 
ad 3.1 mm diam., glabra, seminibus triangularibus, usque 
ad 1.6 mm longis, minute reticulars, hilo punctiformi. 
Distribution and habitat : Known only from one location 

in Brazil. 

Flowering: The one specimen available was in flower 

and fruit in March. 

SPECIMENS EXAMINED 

Brazil, minas gerais: ad Lapoa Santa, 27 Mar. 1864, Warming 
10f>9/2 (c) ; without precise location: Warming (c). 

This rarely collected species is similar to Tripogandra 
diuretica from which it differs in the shape of the leaf 
base, the unique presence of a few uniseriate hairs at the 
junction between adjacent sepals, the flower color, the 
surface texture of the testa and duration (apparently). 

15. Tripogandra palmed (Rose) Woodson, Ann. Missouri 

Bot. Gard. 29:153. 1942. 
Tradcsccvntia palmeri Rose, Contr. U.S. Nat. Herb. 
1:113. 1891. TYPE: Mexico. SONORA: Alamos, 16- 
30 Sept. 1890, Palmer 737 (Holotype, us!; isotypes, 

gh!, ny!). 
DescaJitaria palmeri (Rose) Bruckner, Notizbl. Bot. 

Gart. Berlin-Dahlem 10:56. 1927. 

Illustration: Matuda, Anales Inst. Biol. Nac. Mexico 
26:374. 1956 ('1955'). The magnification is given as X i/ 2 
but it is more nearly XI. 

Plants annual; stems erect, to 45 cm tall, to 4.0 mm in 
diam. when dry, unbranched to branched at every node 
(6 branches) ; internodes to 11.7 cm. long, green, glabrous 
to pilose. Leaves elliptic, narrowly ovate, or ovate ; blades 
0.7-9.5 cm long, 0.2-4.0 cm wide, cuneate, rounded or trun- 
cate at the base, dorsal surface glabrous or pilose, ventral 
surface variously glabrous, glabrous with a few uniseriate 
hairs along the base of the midvein, or pilose, margin 



11)75] Tripogandra — Handlos 289 



ciliate, apex acute; sheaths 1.0-9.5 mm long-, 0.6-9.5 mm in 
diam. when dry, villous at the orifice, with a line of hairs 
extending down the side opposite the blade, otherwise 
glabrous or pilose. Inflorescences borne terminally and in 
the axils of the upper leaves; peduncles to 5.4 cm long, or 
so reduced that cincinni appear sessile, green, proximally 
glabrous or pilose, hairs uniseriate, distally variously 
glabrous, with a few scattered hairs, or pilose, hairs capi- 
tate; cincinni with 2-21 buds, flowers and/or fruits; pedi- 
cels 1.5-8.0 mm long at anthesis, 0.3-0.6 mm in diam., 
green, variously glabrous, with a few scattered capitate 
hairs, or pilose, erect at anthesis and in fruit; bracts at 
the base of the pedicels with an entire margin, glabrous 
or with a few scattered capitate hairs. Flowers white ; 
sepals cymbiform, elliptic to ovate, 1.5-4.1 mm long, 1.0- 
2.3 mm wide, green with the tip darker, pilose or with a 
few scattered capitate hairs, margin entire and hyaline, 
apex acute or obtuse ; petals elliptic, ovate, or obovate, 2.0- 
4.5 mm long, ca. 1.0-5.0 mm wide, cuneate at the base, apex 
obtuse or rounded ; stamens 3, opposite the sepals, shorter 
than the staminodes, filaments 0.5-1.2 mm long, colorless 
with a few (1-6) uniseriate hairs on the upper half of the 
dorsal side (Fig. 13), anthers 0.3-1.0 mm long, 0.2-0.6 mm 
wide, white, basifixed, with white anther sacs, pollen white; 
staminodes 3, opposite the petals, longer, slightly epipeta- 

lous, filaments 1.5-3.4 mm long, white, glabrous, slightly 
bent in the middle (Fig. 21) or S-shaped above the middle 
(Fig. 22), anthers 0.3-1.0 mm long, 0.4-0.9 mm wide, 
broadly basifixed, not versatile, connective yellow, incon- 
spicuous, broadened with anther sacs parallel, or C-shaped 
with the anther sacs divergent, pollen white or yellow; 
ovary 0.5-1.0 mm long, 0.4-0.9 mm in diam., white, 
glabrous, style 0.2-0.4 mm long, stigma simple. Capsule 
globose, 2.0-3.3 mm long, brown or green, glabrous; seeds 
one per locule, ovate with the broad end and two sides 
rolled inward (involute) which results in concavities on 
three sides due to the inrolling, these concavities alter- 
nating with three convexities where the edge is not rolled 



290 



Rhodora [ Vo1 - 77 



(turned), involute margins 



em 



(Fig. 72), testa black, areolate. 



hilum linear (Fig. 73). 

Chromosome number: w=16. 

Distribution and habitat: Mexico, in the states of So- 
nora, Sinaloa, Jalisco, Michoacan, Guerrero, and Puebla; 
at low elevations, 50 to ca. 1400 m. The plants grow in 
relatively dry areas with seasonal rainfall; the vegetation 
is characterized as thorn scrub; candelabra cacti usually 
are abundant. The soil in which these plants grow is 
brown or black loam with abundant humus but in dry 
stream beds it is loose due to the presence of sand and 

gravel. 

Flowering: Plants flower in August, September and 
October. The flowers commonly open about 8:00 AM and 
close about noon. 



I 



REPRESENTATIVE SPECIMENS 

Mexico, sonora: Chorijoa, Rio Mayo, 22 Aug. 1935, Gentry 1598 
(ariz, f). sinaloa: El monte, Los Labrados, 50 m., 15 Oct. 192fi, 
Mexia 927 (UC, us). Jalisco: near Bolanos, Sept. 1897, Rose 2891 
(GH, K, NY, us), michoacan: Apatzin^an, Dist. Apatzin^an, 300 m., 
19 Aug. 1938, Hinton et al 12051 (GH, k, ny). Mexico: Cerro de 
Los Capulines, Palmar Chico, 1100 m., 26 Aug. 1954, Matiula 31337 
(mexu). Guerrero: Coyuca-Chacamerito, Dist. Coyuca, 19 Sept. 
1934, Hinton et al 6620 (GH, K, mich, ny, us). PUEBLA: km. 232.5 
of Hwy. 190, ca. 4 km. west of Rio Atoyac, 1370 (1040)m., 9 Sept. 
1967, Handlos 403 (bh). 

Tripogandra palmeri is unique because the seeds are 
borne one per locule; at anthesis there are two ovules in 
each locule but the lower one always aborts. The seeds are 
further unique in that the margins are involute on three 
sides, rolling- up in such a fashion that they nearly cover 
the protuberant embryotega. 

An interesting pattern of morphological variation can 
be seen in the 20 collections I have examined. All 15 col- 
lections made north of Guerrero or north of 18° 30' N 
latitude have narrowly ovate, nearly glabrous leaves, few 
flowers per inflorescence, a narrow connective, parallel 



11>75 ] Tripogandra — Handlos 291 



anther sacs and epipetalous filaments which are bent 
slightly in the middle. The five collections made south of 
18° 30' N in the southern drainage area of the Rio Balsas- 
Rio Mexcala have larger, more nearly ovate leaves which 



comm 



(though some plants 



mi 



more 



tive is C-shaped, anther sacs are divergent, and the epipe- 
talous filaments are S-shaped. Both of these forms were col- 
lected in the drainage basin of the Rio Balsas. As the plants 
are annuals and self-pollination is probably the most com- 



mon 



be expected to be small. Since the two morphological types 
occur in different geographical areas they might be recog- 
nized as subspecies. Because so few collections have been 
made, a more complex pattern of variation may emerge 
in the future and I decline, therefore, to introduce a new 



name 



lm 



number of collections is rather extensive, 1140 and 360 
kilometers for the northern and southern forms respec- 
tively. Further studies and collections should be made 
throughout the range of this species and especially in the 
Ckierrero-Michoacan border area in the region of Presa 
del Tnfiernillo and to the east which is the area of pre- 
sumed contact between the two forms. 

16. Tripogandra purpurascens (Schauer) Handlos, Baileya 

17:33. 1970. 

Tradescantia purpurascens Schauer in Nees ab Esen- 
beck and Schauer, Linnaea 19:700. 1847. holo- 
type: Mejico, Asckenborn 493 (b!). 

Plants annual; stem erect, to 73 cm tall, branched or un- 
branched, lower portion sometimes decumbent, then root- 
ing at the nodes; internodes to 12.4 cm long, glabrous or 
with scattered capitate hairs, a line of uniseriate hairs 
always present. Leaves narrowly ovate; blades to 8.0 
long, to 2.7 cm wide, complicate, with base rounded or 



cm 



sometimes 



(rarely with a 



292 



Rhodora t Vo1 - 77 



few hairs dorsally), margin ciliate, apex acute; sheaths 



mm lone:, to 8.0 mm 



long hairs at the orifice, a line of uniseriate hairs extending 
down the side opposite the blade, otherwise glabrous or 

lone:. Inflorescences 



mm 



of 1-5 double cincinni borne terminally and in the axils of 
the upper leaves; peduncles to 14.1 cm long, always with 
scattered capitate hairs and with or without 1-2 lines of 
uniseriate hairs extending down the sides; double cincinni 



/ 



mm long, renexect in irun, puuae or wiui »wuci CU *,«*,*«.«- 
hairs (Fig. 1) ; bracts at the base of each pedicel pilose 
or with scattered capitate hairs, margin partly, wholly, 
or not at all ciliate. Flowers pink or rarely white; 
sepals ovate-cymbiform, to 5.0 mm long, to 2.8 mm wide, 
green or with a red tip and/or base and/or margin, pilose 
or with scattered capitate hairs (Fig. 1) to 1 mm long, 
margin hyaline, apex more or less acute; petals obovate, 
to 6.0 (-8.6) mm long, to 4.8 mm wide, base cuneate, apex 



acuminate 



ment 



middle, white on the ends, bearing a tuft of moniliform 



m 



0.4-1.1 mm long, 0.4-1.2 mm wide, white with a purple 
line around the open sac, basifixed, anther sacs more or 

less oarallel, pollen white; inner whorl of stamens longer, 



mm 



giabrous, bent in an open S-shape and slightly expanded 
in the upper portion (Fig. 18), anthers 0.5-1.1 



mm 



mm 



anther sacs divergent or nearly parallel through bending 



m 



mm in diam 



long, stigma simple, capitellate or capitate. Capsule obo- 



mm loner, 1.8-4.0 mm in diam 



brown, glabrous; seeds two per locule, triangular, 1.2-1.8 
(-2.1) mm long, brown, testa ribbed areolate (Figs. 50, 
51, 54, 55), hilum punctiform (Figs. 51, 55). 



1975] Tripogandra — Handlos 298 



In the past, plants of this species were identified as 
THpogandra disgrega. As noted earlier, the holotype for 
T. disgrega is unmistakeable so another name must be 
applied to these plants. The next available name is Tru- 
deseantia purpurascens Schauer. The description accords 
well with Aschenborn 493 in the Berlin herbarium and 
because I have seen no other specimen I consider this to 
be the holotype. The description does not mention the 

; or seeds but dissection of the flowers proved the 
n to be THpogandra. This specimen was seen by 
C. B. Clarke, annotated by him, and is cited apparently 
as "Amberbeau 403", under his description of Tradescantia 
disgrega. Several specimens of T. purpurascens were listed 
bv him under T. amplexicaulis . He apparently did not 
have clear concepts of the species T. amplexicavlis and T. 
disgrega. He does not include the name T. purpurascens 
in his list of synonyms for T. disgrega or any other species. 
This omission is surprising considering the number of 



amen 
pecim 



manuscript nam 



the 



various species he dealt with. 

The separation of and differences between Tripogandra 
disgrega and T. pnrpurasoens are discussed under the 
former species. 

Tripogandra purpurascens is found in every country 
from Mexico to Panama, is absent from most of South 
America and occurs again in southern Bolivia and northern 
Argentina. This outlier is morphologically recognizeable 
by the presence of scattered capitate hairs on all the inter- 
nodes. I recognize these plants as T. purpurascens sub- 
species australis. In all other respects it seems to be 
identical with the typical element of the species. 



16a. Tripogandra purpurascens (Schauer) Handlos subsp. 

purpurascens. 

Tradescantia minata Clarke in DC, Monographiae 
Phanerogamarum 3:307. 1881. type: Mexico. 
Uhde 141a (Holotype, b!; isotype, l!). 



294 Rhodora [Vol. 77 



Trades cantia parvula Brandegee, Univ. Calif. Publ. 
Bot. 6:51. 1914. TYPE: Mexico. PUEBLA: Boca del 
Monte, Oct. 1913, Purpus 6486 (Holotype, uc!; iso- 
types, bm!, f!, gh!, mo!, ny!). 

Descontaria minuta (Clarke) Bruckner, Notizbl. Bot. 
Gart. Berlin-Dahlem 10:56. 1927. 

Tripogavdra minuta (Clarke) Woodson, Ann. Mis- 
souri Bot. Gard. 29:153. 1942. 

Lower internodes glabrous except for a line of uniseriate 
hairs extending down one side, rarely the upper internodes 
with scattered capitate hairs. 

Chromosome number: t? = 16. 

Vernacular name: matlale fide Kerber, Mexico. 

Distribution and habitat: Mexico, Guatemala, Honduras, 
El Salvador, Nicaragua, Costa Rica, Panama; in seasonal 
stream beds or as a weed in cultivated fields at elevations 
from 990 to 2500 meters. 

Flowering: In Mexico flowering occurs from August to 
December while in Guatemala it occurs from September 
to January and occasionally in April and June. In the 
remainder of Central America flowering plants mav be 
found from September to November in Honduras and El 
Salvador, December to February in Nicaragua, July to 
December in Costa Rica, and December to March in 
Panama. 

Flowers open about 10:00 AM and close about 12:45 
PM. 



REPRESENTATIVE SPECIMENS 



Mexico, chihuahua: base of Sierra Mad re, 3 Oct. 1888, Pringle 
1680 (BM, BR, L, M, UC). SINALOA: km. 1184 of Hwy. 40, 1.7 km. 
above Potrerillos, 1600 m., 15 Oct. 1967, Handlos 464 (bh). DURANGO: 
at tbe city of Duranpo and vicinity, 6297 ft., Apr.-Nov. 1896, Palmer 
646 (bm, c, F, GH, MO, NY, us), zacatecas: Sierra de los Morones, 
near Plateado, Sept 1897, Rose 2713 (ny, us), san luis potosi: 
Alvarez, 5-10 Sept. 1902, Palmer 139 (gh, mo, ny, us), nayarit: 
vicinity of Jalisco, 10 Nov. 1925, Ferris 5893 (DS, us). JALISCO: 

about 11 miles southeast of Lagos de Moreno, near highway to Leon, 



1975] Tripogandra — Handlos 295 



1900 m., 7 Sept. 1952, McVaugh 12829 (mich). hidalgo: lower 
slopes of Mt. Lena, west of Lena Station (FCNM), Mun. Nopala, 
Dist. Huichapan, 2500 m., 10 Oct. 1946, Moore, Jr. 1441 (bh, Gh). 
Veracruz: La Luz pr. Cordoba, 7 Oct. 1882, Kerber 90 (bm, c, goet, 
M, us), michoacan: sparsely to densely vegetated slopes of lava 
flow east of San Juan Nuevo, ca. 8 km. south of Uruapan, 6100 ft., 
11-15 Oct. 1961, King & Soderstrom 4719 (MICH, NY, uc, us). Mexico: 
Ixtapan, Dist. Temascaltepec, 1000 m., 19 Oct. 1932, Hinton 2231 
(F, GH, MO, NY, US). DISTRITO federal: Mt. Guadalupe pres Mexico, 

24 Aug. 1865-66, Bourgcau 888 (br, c, gh, k, l, m, ny, us), morelos: 
Las Guacamayas, Cuernavaca, 1800 m., 5 Aug. 1966, Rcbollcdo V. 
(MICH, MSC). puebla: Laguna San Raltasar, vicinity of Puebla, 
2135 m., 20 Sept. 1906, Arsene 327 (us). GUERRERO: Pilas, Dist. 
Mina, 1000 m., 24 Sept. 1937, Hinton 10712 (BR, GH, NY, pom, us), 
oaxaca: Santiago Huitzo, 5500 ft., 22 Oct. 1894, Smith 231 (gh). 
Chiapas: Milpa on the north edge of San Cristobal las Casas, Muni- 
cipio of San Cristobal, 7100 ft., 25 Sept. 1965, Breedlove 12430 (l)S, 
F, MICH). Guatemala. huehUETENANGO: 3 km. south of Huehue- 
tenango, 1800 m., 30 Nov. 1962, Williams, Molina R. A Williams 
22117 (f, ny). quiche: mts. east of Quiche, 2020 m., 20 Nov.-4 Dec. 
1940, Grant 656 (f, GH p.p., MICH p.p.). SAN MARCOS: San Marcos, 

2000 m., 17 June 1882. Lehman*, 1609 (bm, us), quezaltenango: 

Quezaltenango, Sept. 1876, Bernoulli & Cario 781 (goet). SOLOLA: 
mountain slopes above Lake Atitlan, about 3-5 km. west of Pana- 
jachel, 2100 m., 6-7 Dec. 1963, Williams, Molina R. & Williams 25378 
(F, ny). chimaltenango: near Finca La Alameda, near Chimal- 
tenango, 1830 m., 7 Dec. 1938, Standley 59029 (f). Guatemala: 

Finca Rretana, road between Guatemala and Fiscal, 1200 m., 12 
Dec. 1938, Standley 59667 (f). JALAPA: Laguna de Ayarza, 8000 
ft, Sept. 1892, Heyde & Lux 3885 (gh, us), chiquimula: Volcan 
Quezaltepeque, 3-4 miles northeast of Quezaltepeque, 1500-2000 m., 
8 Nov. 1939, Steyermark 31502 (f). santa rosa: Laguna Los Pinos, 
below Cerro Redondo, 25 Oct. 1942, Steyermark 52168 (f, mo). 
Honduras, morazan: Guamiles sobre las faldas noroeste de la Mt. 
Uyuca, cerca de Las Flores, drainage of the Rio Ye^uare, 1600 m., 
20 Oct. 1948, Molina R. 1273 (F, GH, mo). El Salvador. LA libertad: 
rim of Volcan San Salvador, 1800 m., 22 Sept. 1946, Williams & 
Molina R. 10622 (GH, MICH, mo). Nicaragua, JINOTEGA: road to La 
Fundadora, entering at km. 142 from Managua, region of Santa 
Maria de Ostuma, 1400 m., 7 Dec. 1958, Hawkes, Hjerting & Lester 
2197 (C, K). MATAGALPA: road to La Fundadora, cloud forest area 
north of Sta. Maria de Ostuma, Cordillera Central de Nicaragua, 
1300-1500 m., Feb. 1963, Williams, Molina R. & Williams 24949 
(F, ny). Costa Rica, alajuela: Clairiere au rancho de 1'Achiotc 
(Poas), 2200 m., Nov. 1896, Tonduz 10765 (br, us), san JOSE: 



296 



Rhodora [Vol. 77 



a San Jose, 1135 m., Oct. 1890, Tonduz 3047 (br, m, us), cartago: 
Cartaffo, 4250 ft., Nov. 1887, Cooper 5962 (gh, ny, us). Panama. 
CHlRiQUi: Alto Lino, vicinity of El Boquete, 990 m., 3 Feb.-15 Mar. 
1938, Maurice 743 (us). 

The species Tradescantia minuta has confused various 
authors — Matuda (1956) for example, includes specimens 
of Leptorhoeo and Murdannia in his circumscription of it. 
Clarke's original description states that the staminal fila- 
ments are glabrous. One assumes from this that all six 
are glabrous and if one studies the dissected flower present 
on the holotype (Wide 141a), this seems correct. The 



blem arises from 
immature one an 



flower is 



yet developed. An examination of flowers well past an- 
thesis with nearly mature capsules reveals that the short 
filaments of the outer whorl are bearded and the longer 
filaments of the inner whorl are glabrous. The remainder 
of the description seems correct. Wide 141a seems to repre- 
sent a very reduced (depauperate) form of Tripogandra 
purpurascens subsp. purpuras c ens. The corrected obser- 
vation of the presence of bearded filaments leaves no 
character by which to distinguish this material as a 
different species. 

As Clarke erred in describing his new species with 
glabrous filaments, so T. S. Brandegee erred in describing 
the six filaments of his new species, Tradescantia parvula, 
as bearded. Again, a closer examination of the specimens 
reveals that the three short filaments are bearded but the 
three longer ones are glabrous. Tradescantia parvula 
represents another depauperate form of Tripogandra pur- 
purascens subsp. purpurascens. Plants which have a 
strictly annual habit seem to be able to flower and produce 
seeds when of a very small size and with very few leaves. 
On the other hand some plants seem capable of producing 
long stems. In Mexico, plants of Handles 160 flowered at 
a height of 12.5 cm, but when seeds of these plants were 
grown in the greenhouse in Ithaca, N.Y., the seedlings 
reached a height of over a meter before flowering. 



1975] Tripogandra — Handlos 297 



Throughout Mexico this subspecies is relatively uniform 
in size considering the plasticity of annuals. The presence 
of one or two lines of uniseriate hairs on the peduncle is 
a sporadic character over much of Mexico but in southern 
Mexico and in Central America it becomes more common 
and is used as a key character by Standley and Steyermark 
in their Flora of Guatemala. There is still variability — 
some plants lack the line — but the frequency of occur- 
rence is much higher than in Mexico north of Chiapas. 

The colonizing and weedy propensities of Trli)ogandra 
purpurascens subsp. purpurascens should be noted. Plants 
of this subspecies probably existed in stream beds and 
naturally disturbed areas such as those around volcanoes 
prior to man's invasion of the ecosystem. Often I have 
seen T. pvrpurascens subsp. purpurascens in seasonal 
stream beds and on August 19, 1965, on a trip to the 
volcano of Paricutin, I found plants of subsp. purpur- 
ascens growing within ten meters of the edge of a lava 
flow. The area had been covered with a layer of volcanic 
ash but the plants were growing, flowering and producing 
seeds under these conditions. Volcanic activity has oc- 
curred extensively and for a long period of time in Mexico. 
Clausen (1959) has designated the central, volcanically 
active area of Mexico as the Trans-Mexican Volcanic Belt. 
The weedy habit in this species probably could have existed 
before the advent of man and his disturbances of the 
natural vegetation but man must be credited for opening 
additional areas for colonization by this species and allow- 
ing it to become the most common species of Tiipogaudia 
in Mexico. 



16b. Tripogandra purpurascens (Schauer) Handlos subsp. 

australis Handlos, subsp. nov. TYPE: Bolivia. Toldos 
bei Bermejo, 1850 m., 26 Nov. 1903, Fiebrig 2221 
(Holotype, GH ! ; isotypes, k!, l!, m!). 



Illustrations: Bacigalupo, Darwiniana 13:402. 1967, 
Figs. 2, 5a-g, as T. disgrega. 



298 



Rhodora [Vol. 77 



Internodia pilis capitatis, dispersis pubescentia. In 
Bolivia australi et Argentina boreali indigena. 

Internodes all with scattered capitate hairs and a line 

of uniseriate hairs. 

Distribution and habitat: southern Bolivia and northern 
Argentina; in wet wooded areas or as a weed in disturbed 
places at elevations from 450 to 1850 meters. 

Flowering: In January, February and March in Argen- 
tina. 



REPRESENTATIVE SPECIMENS 

Bolivia. Fiebrig 2221 (Gir, K, L, m). Argentina. SALTA: Pept. 
Gen. M. Guemes, camino de La Cornisa, between Jujuy and Salta, 
at km. 1648.9, 1480 m., 18 Mar. 1966, Hawkes, Hjerting <C- Rahn 
3909 (c). tucuman: Cuesta del Garabatal, Sierra de Ende, Jan. 
1874, Lorentz <C- Hieronymus 875 (cord, e, f, goet). 

17. Tripogandra saxicola (Greenman) Woodson, Ann. Mis- 
souri Bot. Gard. 29:154. 1942. 
Tradescantia saxicola Greenman, Proc. Amer. Acad. 
Arts 39:70. 1903. TYPE: Mexico. GUERRERO: Iguala 
Canon, 14 Sept. 1900, Pringle 9270 (Holotype, gh!; 

isotype, vt!). 
Descantaria saxicola (Greenman) Bruckner, Notizbl. 

Bot. Gart. Berlin-Dahlem 10:56. 1927. 

Plants annual, erect or with the lower portion decumbent, 
rooting at lower nodes; stems to 31 cm long, unbranched 
to branched at every node; internodes 0.1-7.4 cm long, 
green or purplish-tinged at the base of the plant, a band 
of uniseriate hairs extending from the sheath above to the 
node below, otherwise variously glabrous to pilose. Leaves 
narrowly ovate to ovate; blades 1.0-7.4 cm long, 0.1-2.0 
cm wide, glabrous dorsally, ventrally variously glabrous, 
pilose along the midvein, or pilose over the entire surface, 
base cuneate or truncate, margin ciliate; sheaths 1.0-5.0 
mm long, 1.2-5.0 mm in diam. when dry, villous at the orifice, 
a line of uniseriate hairs extending down the side opposite 
the blade, otherwise glabrous or with a few scattered uni- 
seriate hairs. Inflorescences few, borne terminally and in 



197r> ] Tripogandra — Handlos 299 



the upper leaf axils; peduncles to 5.9 cm long, or so 
reduced that the double cincinni appear sessile, glabrous 
except for one or two lines of uniseriate hairs extending 
down the side; double cincinni with 1-10 flowers; pedicels 
2.2-5.5 mm long, 0.3-0.6 mm in diam., pilose to densely 
pilose (Fig. 2), reflexed in fruit; bracts at the base of 
each pedicel usually ciliate. Flowers white; sepals ovate- 
cymbiform, 2.0-4.6 mm long, 1.3-2.4 mm wide, green, 
sparingly pilose to densely pilose, apex acute to obtuse, 
margin entire, hyaline; petals elliptic-ovate, ca. 2.5-5.8 
mm long, ca. 2.0-4.5 mm wide, tapering at the base, apex 
obtuse ; stamens 6, in two whorls, those of the outer whorl 
shorter, filaments 1.0-1.6 mm long, glabrous (Fig. 12), 
anthers 0.6-1.0 mm long, 0.3-0.7 mm wide, basifixed, versa- 
tile with parallel white anther sacs which are longer than 
the gibbous white connective, pollen white; stamens of the 
inner whorl longer, slightly epipetalous, filaments 2.0-4.5 
mm long, bearded on the dorsal distal surface, with white 
moniliform hairs (Fig. 20), connective U-shaped, 0.3-0.7 
mm long, 0.3-0.9 mm wide, yellow, dorsifixed, versatile, 
with yellow divergent anther sacs, pollen yellow; ovary 
0.7-1.1 mm long, 0.6-1.1 mm in ( 

long, stigma capitellate. Capsule glabrous, 



lam 



mm 



mm 



triangular or triangular with a notch in one end, with 2-3 



ming 



brown, surface areolate (Fig. 56), hilum punctiform (Fig. 



57) to elliptical. 

Chromosome number: n=21. 

Distribution and habitat: in the states of Guerrero, 
Morelos, and Puebla, Mexico; in thin soil in rocky areas 



Flowering: 



m 



PM 



5:00 PM. Therefore, this species appears to be unique 
among the Mexican Tripogandra species because it is the 
only one in which the flowers open in the afternoon; all 
other species flower in the morning even though blossoms 
may not close until afternoon. 



300 



Rhodora t VoL 7T 



BKPRKSENTATIVE SPECIMENS 



Mexico. MORELOS: Yautepec, near Cuernavaca, 22 Oct. 1902, 
Pringle (vr). puebla: km. 216 on road to Oaxaca, Hwy 190 about 
16 km. SE of Izucar de Matamoros, 30 Au ff . 1965, Handle* 21, 
(bh). Guerrero: Canon del Mano, along railroad tracks north ot 
Iguala, ca. 3 km. north of El Naranjo, ca. 840 m., 13 Sept. 1 .)<>., 



Handles 418 (bh). 



may 



growth cvcle of Tripogandra saxicola. In Mexico this 
species acts as an annual. All collections seem to be of the 
current season's growth judging from the presence of 
juvenile leaves at the base of the plant and the absence 
of old trailing stems and interconnections between plants. 
In the greenhouse, however, plants of Handlos 215 have 
functioned as perennials. The plants lose most of then- 
leaves during the winter, though flowering continues, and 
some of the stems remain green. In May and June, vege- 
tative shoots are produced, the plants grow vigorously, 
rooting at the nodes, and vegetative reproduction is very 
easy. However, not all collections react in this fashion. 



Ha 



and died completely after flowering and producing seeds 
in 1967 and 1968. This behavior seems significant as it 
indicates that both the annual and perennial habit may 
exist within some species. With the extinction of all the 
perennial plants the species would appear strictly annual. 
The presence of annual and perennial populations also 
occurs in Tripogandra semdata and may indicate that 
the annual habit has arisen independently several times 
within the genus and that some annual species may be of 

recent origin. 

The collections Handlos 198 and 419 merit mention be- 
cause these plants were collected in the same area in two 
different years. In 1965, when Handlos 198 was collected, 
the area was covered with large shrubs and small trees 
and had received enough rain so the ground was moist 
and water dripped from the ledges. Plants reached a 
maximum height of 31 centimeters and were found only 



1975] Tripogandra — Handles 301 



after crawling under and through the bushes. Two years 
later I found the whole area cut over, apparently for fire- 
wood, with very little brush left. It was then possible to 
walk and climb over the rocks with little interference 

the remaining woody vegetation. The plants col- 



om 



H 



was exceedingly dry, and the few individuals which were 



maximum 



This information not only gives an impression of how 

occur in an area from year to year 



much variation may 



mpse 



mi 



there is a reduction in population size, and lastly, extinc- 



the rocky ledges and weedy 



from 
com- 



lum 



18. Tripogandra serrulata (Vahl) Handlos, Baileya 17:33. 

1970. 
Commelina serrulata Vahl, Eclogae Americanae 2:4. 



1798. holotype: Ryan (c!). 

? Tradescantia elongata Meyer, Primitae Florae Es- 
sequeboensis . . . 146. 1818. holotype: apparently 

lost. 

Tradescantia congesta Martens & Galeotti, Bull. Acad. 
Bruxelles 4(2):377. 1842. HOLOTYPE: Mexico. 
VERACRUZ: Mirador, 3000 ft., June-Oct. 1840, Gale- 
otti 4949 (br!). 

Tradescantia balbisii Kunth, Enumeratio Plantarum 
4:97. 1843. TYPE: Portorico, 1834, Balbis (Holo- 
type, b!; isotype, b!). 

Tradescantia schlechtendalii Kunth, Enumeratio Plan- 
tarum 4:94. 1843. type: Mexico. Hacienda de la 
Laguna, Oct. 1828, Schiede 972 (Holotype, b!; iso- 
type, hal!). 

Tradescantia schomburgkiana Kunth, Enumeratio 
Plantarum 4 :663. 1843. type: Guiana Angl., 1842, 
Schombitrgk 189 (Holotype, b!; isotype, k!). 



302 



Rhodora [Vol. 77 



Tradescantia guianensis Miquel, Linnaea 18:374. 1845 
('1844'). TYPE: Surinam. Focke 400 (Holotype, 
u!; isotype, gh!). 

Descantaria tbalbisii Hasskarl ex Clarke in DC, 
Monographiae Phanerogamarum 3:303. 1881, nom. 

nud. pro syn. 

Descantaria schlechtendalii Hasskarl ex Clarke in 
DC, Monographiae Phanerogamarum 3:304. 1881, 
nom. nud. pro syn. 

Ueterachtia gaudicJuiudiana Hasskarl ex Clarke in 
DC, Monographiae Phanerogamarum 3 :303. 1881, 
nom. nwd. pro syn. 

Tradescantia cumanensis (3 glabrior Clarke in DC, 
Monographiae Phanerogamarum 3:306. 1881. 
TYPE: Nicaragua. Chontales, 1867-8, Tate 452 
(Holotype, k!; isotype, BM!). 

Tradescantia elongata ft schlechtendalii (Kunth) 
Clarke in DC, Monographiae Phanerogamarum 
3:304. 1881. 

^.Descantaria elongata (Meyer) Bruckner, Notizbl. 
Bot. Gart. Berlin-Dahlem 10:56. 1927. 

tTHpogandra elongata (Meyer) Woodson, Ann. Mis- 
souri Bot. Gard. 29:152. 1942. 

Tripogandra cumanensis f. glabrior (Clarke) Stand- 
ley & Steyermark, Fieldiana : Bot 24(3) :36. 1952. 



Illustrations: Standley and Steyermark, Fieldiana: Bot. 
24(3) :39, 1952, Fig. 9, a line drawing of a flower and 
upper part of the plant. The position of the stamens is 
incorrectly shown in the open flower and misplaced in the 
flower drawing. 

Plants perennial, decumbent at the base, rooting at 
nodes, flowering stems erect; stems to 1.14 m long, 
branching irregularly; internodes to 14.1 cm long, glab- 
rous except for a line of uniseriate hairs extending 
down the side from the sheath above (Fig. 4). Leaves 
narrowly ovate, occasionally ovate; blades to 13.2 cm long, 



1975] Tripogandra — Handlos 303 



to 2.7 cm wide, glabrous to pilose dorsally, rarely with a 
line of hairs along the midvein, glabrous to sparingly pilose 
ventrally, often with a line of uniseriate hairs along the 
midvein, base oblique, margin ciliate, apex acute; sheaths 
to 2.1 cm long, to 1.2 cm in diam., villous at the orifice, a 
line of uniseriate hairs extending down the side opposite 
the blade (Fig. 4), otherwise variously glabrous or the 
uppermost occasionally with scattered uniseriate hairs or 
pilose. Inflorescences composed of 1-13 double cincinni 
borne terminally and in the axils of the upper 1-4 leaves ; 
peduncles 0.4-5.1 (-6.7) cm long, glabrous or with 1-2 lines 
of uniseriate hairs extending down the sides; double cin- 
cinni with up to 17 buds, flowers and/or fruits; pedicels 
0.7-5.0 mm long, 0.5-0.7 mm in diam., reflexed in fruit, 
glabrous to sparingly pilose (Fig. 3), hairs capitate; bracts 
at the base of each pedicel glabrous, margin entire, ciliate, 
or with a few scattered hairs. Flowers white or pink 
(Fig. 16) ; sepals ovate, 2.5-4.6 mm long, 1.3-2.7 mm wide, 
glabrous to sparingly pilose (Fig. 3), hairs capitate, mar- 
gin hyaline, apex acute to obtuse; petals ovate-elliptic, 
3.5-6.2 mm long, 2.3-4.3 mm wide, base cuneate, apex 
rounded; stamens 6 in two whorls, the outer shorter, fila- 
ments 1.0-1.5 mm long, white, glabrous or with a few (1-3) 
moniliform hairs, anthers 0.5-1.0 mm long, 0.4-1.0 mm 
wide, anther sacs parallel, connective inconspicuous, pollen 
white; stamens of the inner whorl longer, filaments 2.7- 
4.5 mm long, white, bent in an open S-shape, bearded with 
moniliform hairs in the upper half (Fig. 19), anthers 
0.4-0.8 mm lon#, 0.5-1.0 mm wide, basifixed, connective 
elongate and narrow, anther sacs divergent, yellow, pollen 
yellow; ovary 0.6-1.3 mm long, 0.5-1.0 mm in diam., white, 
glabrous, style 0.1-0.3 mm long, stigma capitate, weakly 
3-lobed. Capsule elliptical, 2.0-2.8 mm long, 1.5-2.5 mm in 
diam., light brown, glabrous; seeds usually 2 per locule, 
rounded triangular, 0.9-1.5 mm long, gray or brownish- 
gray, testa reticulate, reticulations lighter in color (Fig. 
38), hilum punctiform (Fig. 39). 
Chromosome number: n = lQ, 24. 



304 



Rhodora [ VoL 77 



inam 



meters 



Vernacular names: suelda fide Bro. Daniel, Colombia; 
siempre viva fide Archer, Colombia; palm grass (Bar- 
badian) fide Standley, Panama; matalin, shiu fide Martinez- 
Calderon, Mexico; yerva del poyo fide Curtiss, Mexico. 

Distribution and habitat: central Mexico, south to 
Panama, Colombia, Ecuador, Peru, Venezuela, Guyana, 

and the Caribbean islands; from sea level to 
about 1500 m. Most specimens with altitudinal informa- 
tion are from intermediate to low elevations. Two records 
are exceptions. Matuda reports 2500 
tion in Mexico, while Heyde and Lux reported 8000 feet 
for a collection in Guatemala. These reports should be 
investigated further. 

Flowering: in Mexico from January to December; in 
Guatemala from August to January and March, May, and 
July ; in British Honduras from October to February and 
April; in Honduras from November to March and May, 
June, August, and September ; in El Salvador from Decem- 
ber to April and July; in Nicaragua in April and De- 
cember; in Costa Rica in every month except January; in 
Panama in every month of the year; in Colombia in 
February, March, April, May, June, August, October, 
November, and December ; in Ecuador in May and August ; 

and in Peru in April. 

Flowers open in the morning about 9:00 AM and close 

about 2:30 PM. 

REPRESENTATIVE SPECIMENS 

Mexico, san LUIS potosi: Tamazunchale, 250 ft., 5 Aug. 1937, 
Fisher (MO, NY, us), hidalgo: wooded slopes between Calnali and 
Huazalingo, Dipt. Huejutla, 29 May 1947, Moore, Jr. 3016 (bh). 
Veracruz: Orizaba, Mt. Orizaba, 4000 ft., 29 Aug. 1891, Seaton 29 
(F, GH, NY, us), distrito federal: Monte Guadalupe pres Mexico, 
24' Aug. 1865, Bourgeau 888 (gh). morelos: Cuernavaca, 29 Oct. 
1903, Holway 5261 (gh). puebla: near Metlaltoyuca, 800 ft., 31 
Jan. 1898, Goldman 49 (us), oaxaca: Yaveo, Arroyo del Perrico, 
Dist. Choapam, 475 m., 15 Mar. 1938, Mexia 9153 (F, GH, mo, ny, 
U, UC, us). Chiapas: Escuintla, Nov.-Dec. 1937, Matuda 2176 (F, 
GH, Midi, NY, UC). Guatemala, alt a verapaz: Finca Mocca, 3200 
ft, 5 Jan. 1920, Johnson 169 (NY, us), izabal: vicinity of Quirigua, 



ii)75] Tripogandra — Handlos 305 



75-225 m., 15-31 May 1922, Strtndley 24165 (GH, NY, us). SAN 
marcos: river 5 mi. W of Malacatan, 400 m., 20 Nov. 1940, Grant 
566 (F, gh). quezaltenango: Finca Pireneos, below Santa Maria 
de Jesus, 1350-1380 m., 11 Mar. 1939, Standley 68347 (f, mich). 
sacatepequez : Ciudad Vieja, Mar. 1915, Tejada 343 (us). Guate- 
mala: Brefias y seta cerca de Guatemala, 1400 m., July 1921, Tonduz 
669 (us), jalapa: Laguna de Ayarza, 8000 ft., Oct. 1892, Heyde & 
Lux 3882 (GH, K, M, us), zacapa: Gualan, 420 ft., 20 Jan. 1905, 
Deam 404 (gh, mich). RETALHULEU: San Felipe, 13 Jan. 1917, 
Holway 710 (us), suchitepequez: Chojoja p. Mazatenango, Sept. 
1867, Bernoulli 469 (rr, ny). escuintla: Escuintla, 1100 ft., Mar. 
1890, Donnell-Smith 2220 (gh, m, us), santa rosa: Rio Maria 
Linda, 3000 ft., Sept. 1893, Heyde & Lux 6251 (gh, us), san Marcos: 
Cangutz, 1140 m., 31 July 1922, Galas 11 (us). British Honduras. 
Gracie Rock, Sibun River, 15 April 1935, Gentle 1594 (f, gh, mich, 
mo, ny, us). Honduras, santa Barbara: Rio Permejo, 600 ft., Dec. 
1888, Thieme 5532 (gh, us), cortes: in ravine near Lake Yojoa, 
Agna Azul, 630 m., 28 Dec. 1946, Williams & Molina R. 11411 (BH, 
GH, mich, MO, uc). comayagua : Rittenhouse's hacienda near Sigua- 
tepeque, 1050 m., 30 June 1936, Yuncker, Dawson & Youse 5527 
(f, GH, mich, mo, u). atlantida : Ceiba, 26 Sept. 1916, Dyer A94 
(us), yoro: Farm 39 of the Tela Railroad Company, Guaymas 
Dist., 30 m., 2 Feb. 1928, Standley 55489 (us), olancho: a la orilla 
de la quebrada cerca de El Plomo, Valle Catacamas, 300 m., 19 Nov. 
1963, Molina /?. 13290 (g, ny). lempira: faldas de Montana Puca 
cerca de Los Cuabanos, 1300 m., 25 Sept. 1963, Molina R. 12956 
(f, ny). morazan: along Santa Clara Creek, drainage of the Rio 
Yeguare, 850 m., 6 Aug. 1949, Williams A Molina R. 15865 (on, us). 
el paraiso: Montana entre Cifuentes y El Urraco, 900 m., 15 Mar. 
1963, Molina R. 11428 (f, NY, US). El Salvador. aHUAChapan: 
vicinity of Ahuachapan, 800-1000 m., 9-27 Jan. 1922, Standley 19838 
(GH, NY, us). SONSONATE: Finca Chilata, 26, 27 Dec. 1921, Standley 
19311 (gh, ny, us). LA LIBERTAD: vicinity of Santa Tecla, 790-950 
m., 10 Apr. 1922, Standley 23054 (us), san Salvador: San Salvador, 

July 1922, Calderon 913 (gh, mo, ny, us), la paz: Zacatecoluca, 

Mar. 1922, Calderon 303 (gh, ny, us), san VICENTE: vicinity of 
San Vicente, 350-500 m., 2-11 Mar. 1922, Stdndley 21727 (gh, mo, 
NY, us). Nicaragua. JINOTEGA: road to La Fundadora, entering at 
km. 142 from Managua, region of Santa Maria de Ostuma, 1400 m., 
7 Dec. 1958. Hawkes, H jetting & Lester 2198 (c). chontales: 
slopes of Mt. Mombacho, near Grenada, 460 m., 18 Dec. 1940-9 Feb. 

1941, Gran* 787 (f, git, mich). zelaya: vicinity of El Recreo, on 

Rio Mico, ca. 30 m., 23 Apr.-14 May 1949, Standley 19091 (f). 

Costa Rica, ala.tuela: Villa Quesada, Canton San Carlos, 825 m., 

10 Mar. 1940, Smith p2574 (f, mich, mo), san JOSE: vicinity of 



306 



Rhodora [Vol. 77 



El General, 915 m., Feb. 1936, Skuteh 2602 (gh, mich, mo, ny, us). 
CAktago: Angostura, 19 June 1874, Kuntze 2050 (ny). umon: 
Jimenez, Llanos de Santa Clara, 650 ft., Apr. 1894, 1 ) onnell-Smith 
4976 (GH, k, us). Panama, bocas del toro: Bocas del Toro, fi Nov. 
1920, Carleton 71 (GH, ny, us), chiriqui: Boquete, Boquete Dist., 
3800 ft., 17 May 1938, Davidson 665 (f, gh, mo, us), cocle: Lower 
Rio Anton, vicinity of El Valle De Anton, 800-1000 (600) m., 30 
Dec. 1936, Allen 111 (gh, mo, ph). canal zone: ruins of fort, Fort 
San Lorenzo, Fort Sherman Military Reservation, 14 June 1923, 
Maxon & Valentine 7018 (c, gh, us). Panama: Tumba Muerto 
Road, near Panama, 6 Jan. 1924, Standley 29717 (c, us), darien: 
vicinity of Boca de Cupe, ca. 40 m., 5 Oct. 1938, Allen 890 (f, gh, 
MO, NY, us). SAN blas : Perme, 24 Apr. 1933, Cooper III 253 (NY, 
us). Venezuela, carabobo: Represa del acueducto de San Esteban, 
Pto. Cabello, 7 Jan. 1965, Aristeguieta 5421 (ven). aragua: entre 
Guamilas y Rancho Grande, P.N., 850 m., 6 Oct. 1938, Williams 
10382 (ven). federal district: Caracas and vicinity, 3000-3500 ft., 



Bailey 



Miranda: bosque de Los 



Guayabitos, arriba de Baruta, Nov. 1964, Aristeguieta 5395 (ven). 
merida: 3% miles west of city of Merida, 5000 ft., 24 Jan. 1931, 
Reed 288 (us), bolivar: San Jose, Ciudad Bolivar and vicinity, on 
the Orinoco, about 200 ft., 9 Jan. 1921, Bailey & Bailey 811 (bh). 
Colombia, magdalena: Minca road, Santa Marta, 1200 ft., 23 Nov. 

1898-1899, Smith 2280 (BM, BR, E, F, GH, L, MICH, MO, NY, PH, u, uc, 

us, vt, wis), bolivar: Los Hurtados, on Rio Sinu, 40-70 m., 4 Feb. 
1918, Pennell 4153 (ny, us). CHOCO: Anda^oya, 70-100 m., 20-30 
Apr. 1939, Killip 35075 (bm, mo, us), antioquia: vicinity of Medel- 
lin, 10 Mar. 1927, Toro 37 (ny, us), santander: Puerto Wilches 
and vicinity, 100 m., 28 Nov.-2 Dec. 1926, Killip & Smith 14773 (ny). 
valle del cauca : Estero de Congrejo, north shore of Buenaventura 
Bay, near sea level, 3 June 1944, Killip & Cuatrecasas 38730 (f, its). 
CALDAS: Santa Cecilia, Cordillera Occidental, Vertiente Occidental, 
800 m., 16 Feb. 1945, von Sneidern 5181 (us), tolima: "La Trini- 
dad/' Libano, 1000-1200 m., 21-25 Dec. 1917, Pennell 3363 (ny). 
Ecuador, pichincha: entre Santo Domingo y la Hcda. Lelia, Sec- 
tion Occidental, 400-800, 950-1100 m., 11 Aug. 1945, Acosta-Solix 
10937 (F, us), imbabura: entre El Pajon y Cachaco, 600, 740 m., 
30 May-12 June 1949, Acosta-SolU 12728 (f, us). Peru, tumbes: 
a 8 km. al sur de Tumbes, 15-20 m., 24 Apr. 1949, Ferreyra 6002 
(us), loreto: near km. 194 below Divisoria on road from Tingo 
Maria to Aguaytia, Prov. Coronel Portillo, ca. 1400 m., 22 Dec. 1960, 
Moore, Jr., Salazar C. & Smith 8632 (bh). junin: La Merced, 
ca. 700 m., 29 May-4 June 1929, Killip & Smith 23416 (ny, us). 
Guyana, pomeroon district: Mora Landing, Moruka River, 21-23 
Aup. 1922, De La Cruz 1844 (bh, f, gh, mo, ny, us). Surinam. 



1975] Tripogandra — Handlos 307 



Suriname R. near Gansee, 15 Nov. 1933, Lanjouw 1307 (ny, u, us). 

Dominican Republic. Mad re Vieja, Nagua, Prov. Maria Trinidad 

Sanchez, 20 Dec. 1964, Jimenez -1108 (ny). Puerto Rico. San Juan, 

14 km. S on Mil. Road, 2 Mar. 1899, Heller & Heller 662 (F, ny, us). 

Guadeloupe. 1895, Dttss 3619 (f, ny, us). Dominica, prope Wotten 

Waven, 200 m., Dec. 1887, Eggers 690 (br, cord, goet, l, m, uc). 

Martinique. La riviere du Galion (Trinite) et de la riviere du Carbet, 

Oct. 1888, Dues 1024 (ny). St. Vincent. 1000 ft., March 1890, Smith 

& Smith 1660 (bm). Trinidad. North Range, roadside forest, Arima 

Valley Road, 500 m., 24 Mar. 1959, Cawa)n & Simmonds 1180 (ny, 
US). 



om 



me has not 



other species by its gray or gray-brown seeds with lighter 
colored reticulations which produce a cobweb pattern over 
the surface. 

Within Tripogandra serrulata there is variation in leaf 

shape as well as in the vesture of leaves, peduncles, pedicels, 

and sepals. I have not seen any consistency or pattern in 

this variation. This does not of necessitv mean that none 

exists, however, only that my study has not been precise 

enough and that the information available 1 
been complete. 

Many of the specimens associated with this species have 
been identified in the past as Tripogandra cumanensis. 
This seems to be the result of a misidentification of speci- 
mens by Clarke. The name which has priority is Com- 
melina serrulata. Tradescantia cumanensis is cited as a 
synonym of Tripogandra mnltiflora and is discussed under 
that species. 

Vahl's description of Commelina serrulata agrees well 
with the type specimen, but does not mention the stamens 
or the seeds. It would be difficult to know from the descrip- 
tion alone to which species or even genus this specimen 
should be referred. The specimen has nearly 
which have a reticulation matching that on 



mat 



recent taxonomists have called Tripogandra cuma?iensis. 
The stamens follow the diagnostic Tripogandra pattern ; 
they are dimorphic with the shorter ones opposite the 



308 



Rhodora [Vol. 77 



sepals. 



imo 



that Vahl placed this species in the genus Comm 



name 



years but the type specimen is undoubtedly a Tripogandra. 
Kunth (1843) included this species in the genus CommeUtw 
in a section called "Species valde dubiae," indicating that 
he was not certain what the plant was. His decription was 
taken from Vahl and in parts is nearly a word-for-word 
copy, implying that he had probably not seen a specimen. 
The problem of the identity of Tradescantia elongata 
has plagued taxonomists for many years. I have not seen 
the type specimen; it should be a Rodshied collection 
(Stearn and Williams, 1957) preserved at Gottingen but 
was not located among the specimens obtained on loan. 
If T. elongata does actually apply to Tripogandra it can 
only apply to the one taxon which seems to occur in 
the vicinity of the Essequibo River in Guyana. The plants 
from this area are of great interest and should be studied 
further. I have examined sixty-one sheets of specimens 
from this area: all sheets have several inflorescences but 
none of them have fruits in any stage of development. In 
addition I mounted pollen in aniline blue-lactophenol from 
some of the plants identified as T. elongata from through- 
out its range. The pollen does not stain and on that basis 
is considered to have been inviable when fresh. From per- 
sonal experience with T. serndata in Mexico and knowledge 
of the existence of sterile plants within these populations, 
I have treated all these sterile collections as part of the 
species T. serndata, but only after studying and describing 
that species from fertile specimens. The pattern and range 
of variation within the sterile collections falls within that 
of fertile plants of T. serrulata, except for the peduncle 
length of three collections from Guyana. The longest pe- 
duncles were 5.7, G.4, and 6.7 cm. while the longest peduncle 
on fertile T. serrulata was 5.1 cm. I do not consider this 
problem solved. The apparent sterility of these plants 
must be studied in the field to see if it is characteristic 
or whether my sample by some quirk is very biased. 



975] Tripogandra — Handlos 309 



Tripogandra elongata has been interpreted broadly in 
the past and three taxa have been included within it. These 
are T. diuretica of southern Brazil, Bolivia, Paraguay, 
Agentina, and Uruguay, T. montana of Central America, 
and T. elongata sensu stricto of eastern Venezuela, Guyana 
and Surinam. These three taxa are not sympatric in any 
part of their ranges and are morphologically distinguish- 
able. Therefore, T prefer to recognize T. diuretica and 
T. montana, as separate and distinct species, while the 
sterile plants of T. elongata are included within T. ser- 
rulate. 

Kunth (1843) proposed the name Tradescantia schlech- 

tendalii based on Schiede 972, which Schlechtendal had 
misidentified and called Commelina mexicana Presl. PresFs 
original description contains several points indicating that 



the plants were different: 



Stam 



tria. Filamenta erecta fertilia hirsuta, duo pistillo breviora, 
tertium pistillo longius. Antherae . . . apice poro dehis- 
centibus polliniferia . . . Stylus simplex curvatus. Stigma 
eniarg^'natum." The Schiede specimen has white petals, 
six stamens, three longer and three shorter, and a capitate 
stigma. Kunth proceeded, however, to recognize Com- 
melina mexicana Presl as a species of Tradescantia. I 
believe that this name may apply to a species of Com- 
melina though I am not certain of this. In any case it 
certainly is not a species of Tradescantia or Tripogandra, 
on the basis of the description given by Presl. 

Clarke's variety ,6 glabrior of Tradescantia cumanensis 
does not seem to differ from T. congesta. Clarke states 
that the pedicels and sepals are glabrous when, in fact, 
there are a few hairs to be found on the type specimen 
though some structures are glabrous. These two collec- 
tions are certainly conspecific but Clarke considered gla- 
brior to be a variety of T. cumanensis while T. congesta 
was made a synonym of T. elongata. 

Plants of Tripogandra serrulata have been collected from 
several Caribbean islands. All of these collections lack 
seeds except one from Hispaniola. The seeds seem identical 



310 



Rhodora [Vol. 77 



mainland T. serrulata. If this plant repre- 

panH not a rpppnt.lv introduced Triuoaandra. 



me 



which have been called T. elongata, as conspecific with 
T. serrulata. 

19. Tripogandra silvatica Handlos, sp. nov. type : Mexico. 

Veracruz: Montepio, 19 km. al E de Catemaco, 19 
Mar. 1965, Gonzales Quint ero 2239 (Holotype, mich!; 
isotype, msc!). 

Herb a perennis(?); caulis decumbens, usque ad 30 cm 
longus; inter nodiis usque ad 5.5 cm longis, linea unica 
pilorum instructis, aliter glabris. Folia ovata, laminis 
usque ad 2.9 cm longis, usque ad 1.4 cm latis, basi obliquis, 
apice acutis, dorsaliter glabris, ventraliter glabris vel pilis 
dispersis instructis praeter lineam partialem pilorum secus 
costam, margine ciliatis, vaginis usque ad 5.6 mm longis, 
usque ad 3.0 mm diam., orificio villosis vel pilis dispersis 
praeditis, linea unica pilorum instructis, aliter glabris. 
Inflorescentiae terminates, ex 1 (-3) cincinnis duplicibus 
constantes; pedunculi usque ad 2.3 cm longi, glabri vel 
pilis capitatis paucis dispersis instructi ; cincinni duplices 
omnes alabastra, flores, vel fructus usque ad 13 gerentes; 
pedicelli usque ad 4.5 mm longi, maturitate reflexi, gla- 
briusculi ad pilosi, bracteis basi pedicellorum glabris, 
margine erosis, nonnunquam ciliatis. Flores albi ; sepala 
ovata. cucullata, usque ad 3.7 mm longa, usque ad 1.6 mm 
lata, pilis capitatis pilosa, margine hyalina, apice ± ob- 
tusa; petala usque ad 4 mm longa; stamina 6 in verticillis 
duobus, 3 sepalis opposita filamentibus brevibus, usque ad 
1.3 mm longis, glabris vel ?pilis paucis moniliformibus 
praeditis, antheris usque ad 0.5 mm longis, usque ad 0.6 
mm latis, 3 petalis opposita filamentibus longioribus, usque 
ad 2.7 mm longis, glabris, distaliter sigmoideis, antheris 
usque ad 0.7 mm longis, usque ad 0.6 mm latis; ovarium 
usque ad 0.7 mm longum, glabrum, stylo usque ad 0.4 mm 
longo, stigmate capitellato. Capsula obovoidea, usque ad 
2.3 mm longa, usque ad 1.5 mm diam., glabra, basi stipitata, 



1975] Tripogandra — Handlos 311 



seminibus triangularibus, usque ad 1.2 mm longis, reticu- 
latis, hilo punctiformi. 

Distribution and habitat: in the state of Veracruz, 
Mexico, in the area around Colipa-Misantla and Catemaco; 
in wet forest lowlands. 

Flowering: March and April. 

SPECIMENS EXAMINED 

Mexico. Veracruz: Misantla, Mar. 1841, Liebmann (c); inter 
Colipa et Misantla ad Palenque, Mar. 1841, Liebmann (c) ; Colipa, 
Mar. 1841, Liebmann 350 (c); Colipa, Mar. 1841, Liebmdnn (c) ; 
Jalapa, 4000 ft., 3 Apr. 1899, Pringle 7810 (vt). 

Dried specimens of this species resemble Leiandra 
cordifolia superficially — usually bearing only one terminal 
double cincinnus. Dissection of the flower is necessary to 
reveal the dimorphic nature of the stamens. 

20. Tripogandra warmingiana (Seubert) Handlos, comb. 

nov. 

Tradescantia warmingiana Seubert in Warming, 
Vidensk. Meddel. Dansk Naturhist. Foren. Kjoben- 
havn 126. 1872. holotype: Brazil, minas gerais : 
Lagoa Santa, 6 Mar. 1866, Warming 1046 (c!). 

Plants annual (?) ; stem decumbent at the base and root- 
ing at the nodes, to 19 cm long, branched ; internodes to 
5.6 cm long, glabrous except for a line of uniseriate hairs 
extending down the side from the sheath above. Leaves 
narrowly ovate to ovate; blades to 4.4 cm long, to 1.85 cm 
wide, with the base always oblique but narrowed and sub- 
petiolate on the lower part of the plant, glabrous dorsally, 
glabrous ventrally except for a line of hairs at the proximal 
end of the midvein, margin ciliate, apex acute and slightly 
acuminate; sheaths to 4.4 mm long, to 3.1 mm in diam., 
sparingly villous at the orifice, otherwise glabrous except 
for a line of uniseriate hairs extending down the side 
opposite the blade. Inflorescences composed of 1-3 double 
cincinni borne terminally and in the upper leaf axils; 
peduncles to 1.2 cm long, green, glabrous; double cincinni 



312 Rhodora C Vo1 - 77 



with up to 



d/ 



mm long, green, with a few uniseriate hairs near the 
distal end, reflexed in fruit; bracts at the base of each 
pedicel glabrous, margin erose. Flowers white; sepals 
ovate, to 3.4 mm long, to 1.7 mm wide, pilose along the 
midvein, the remainder with scattered uniseriate hairs, 
margin hyaline, apex acute; petals ovate-elliptic (fide 
Warming), to 5.0 mm long, to 2.3 mm wide, apex obtuse 
(fide Warming) ; stamens 6, in two whorls, the outer 
shorter, filaments to 1.2 mm long, glabrous, anthers to 0.7 
mm loner, to 0.5 mm wide, anther sacs parallel; 



stamen 



mm 



of the inner whorl longer, filaments to 3.7 
in an S-shape (fide sketch, presumably by Warming), 
bearded in the lower portion of the upper half with monili- 
form hairs, anthers to 0.5 mm long, to 1.2 mm wide, con- 
nective elongate and anther sacs divergent; ovary to 0.9 



mm loner. 0.8 mm 



mm 



stigma minutely capitellate. Capsule globose, to 3.0 
long, to 2.8 mm in diam., light brown, glabrous; seeds 2 
per locule, rounded-triangular, gray or brown, testa reticu- 
late-foveate (Fig. 61), hilum punctiform. 

Distribution ; known only from three collections at Lagoa 
Santa, Brazil. 



SPECIMENS EXAMINED 



Brazil, minas germs: La^oa Santa, 10 Mar. 1864, Warming 
1070 (778) (c) ; Lagoa Santa, Warming (f, us p.p.). 

I know this species from only three collections. Accord- 
ing to the original description it was found in a wooded 
area above a rocky calcareous site near Lagoa Santa. 
Brazil and flowered from January to March. 

The original description accords well with the type 
specimen. One discrepancy exists concerning the shorter 
stamens. The original description states: "Stamina tria 



minora. . . ." 



stamens 



opposite the sepals and are the outer whorl of stamens. 

C. B. Clarke considered this species as part of his 

Tradescavtia elongata. The habit and size of the plant. 



1975] Tripogandra — Handlos 313 



shape of the laminar base, seeds, flower size and color, and 
stigma show this species to be distinct. 



EXCLUDED SPECIES 

Tripogandra cordi folia (Swartz) Aristeguieta, Bol. Acad. 

Ci. Fis. 25:125. 1965. 

Tradescantia cordifolia Swartz, Nova genera & species 

plantarum sen prodromus ... 57. 1788. 
Leiandra cordifolia (Swartz) Rafinesque, Flora Tellu- 

riana 2:16. 1837 ('1836'). 
Callisia cordifolia (Swartz) Anderson & Woodson, Contr. 

Arnold Arbor. 9:117. 1935. 
Phyodina cordifolia (Swartz) Rohweder, Abh. Aus- 

landsk., Reihe C, Naturwiss. 18:151. 1956. 
T have excluded Tripogandra cordifolia because it has 
six more or less similar, glabrous stamens. The genus 
Phyodina as interpreted by Rohweder is composed of di- 
verse elements which show little affinity. This problem 
has been discussed to some extent by Moore (1963) but 
requires much more study in conjunction with a considera- 
tion of Tradescantia gracilis, T. debilis, and Leptorhoeo 

aii for mis. 

Tripogandra lundellii (Standley) Woodson, Ann. Missouri 

Bot. Gard. 29:153. 1942. 
Tradescantia lundellii Standley, Publ. Field Mus. Nat. 

Hist. Bot. Ser. 22:5. 1940. type: Lundell 7098 (Holo- 

type f!; isotypes, mich!, ny!). 
Gibasis sp. 

Standley's original description of Tradescantia lundellii 
described stamens of two lengths. This species cannot be 
considered one of Tripogandra because at anthesis the 
plants have six essentially similar stamens which are borne 
on flowers in a single cincinnus. 

Tripogandra rosea (Ventenat) Woodson, Ann. Missouri 

Bot. Gard. 29:153. 1942. 
Tradescantia rosea Ventenat, Jard. Cels 24. 1800. holo- 
type: Michaux p; photograph, gh !. 



314 



Rhodora [Vol. 77 



Cuthbertia rosea (Ventenat) Small, Flora of the South- 
eastern United States 237. 1903. 
Phyodina rosea (Ventenat) Rohweder, Abh. Auslandsk., 

Reihe C, Naturwiss. 18:151. 1956. 

Tripogandra rosea lacks the dimorphic stamens which 
characterize Tripogandra sensu stricto. Rohweder's inclu- 
sion of this species in Phyodina should be investigated 
further. 

Tripogandra stenophylla (Brandegee) Matuda, Anales 

Inst. Biol. Univ. Nac. Mexico 26:369. 1956 ('1955'). 

Tradescantia stenophylla Brandegee, Univ. Calif. Publ. 

Bot. 3:377. 1909. TYPE: Purpus 3352 (Holotype, 



uc!; isotype, ny!). 



s 



Tripogandra stenophylla is properly placed in the genu.' 
Tradescantia because this plant has six similar stamens 
and two foliaceous bracts subtending the double cincinnus. 

Tripogandra irarscewicziana (Kunth & Bouche) Woodson, 

Ann. Missouri Bot. Gard. 29:154. 1942. 
Tradescantia ivarszewicziana Kunth & Bouche, Ind. 

Sem. Hort. Berol. 11. 1847. 
Spironema warszeiviczianum (Kunth & Bouche) Bruck- 
ner, Notizbl. Bot. Gart. Berlin-Dahlem 10:56. 1927. 
Phyodina ivarsceicicziana (Kunth & Bouche) Rohweder, 
Abh. Auslandsk., Reihe C, Naturwiss. 18:151. 1956. 
Hadrodemas ivarszewicziana (Kunth & Bouche) Moore, 

Baileya 10:134. 1963 ('1962'). 
Tripogandra ivarsceicicziana must be excluded because 
of its inflorescence structure and a lack of dimorphic 
stamens. Moore (1963) has more fully discussed the place- 
ment of this unusual species. 

Descantaria. laxi flora (Clarke) Bruckner, Notizbl. Bot. 

Gart. Berlin-Dahlem 10:56. 1927. 
Tradescantia laxiflora Clarke in DC, Monographiae 

Phanerogamarum 3:307. 1881. type: Andrieux 51 

(Holotype, k!; isotype, m!). 
Gibasis sp. 



1975] Tripogandra — Handles 315 

This species has a single cincinnus so it cannot be con- 
sidered a species of Tripogandra. 



ACKNOWLEDGMENTS 

The author expresses his appreciation and gratitude to 
Drs. H. E. Moore, Jr., H. T. Stinson, and R. B. Root, 
Special Committee members, for their advice, criticism, 
and assistance. The writer is indebted to Drs. C. H. Uhl, 
N. W. Uhl, G. Eickwort, and W. J. Dress, who have pro- 
vided assistance and encouragement with special aspects 
of this work. Deepest appreciation is due Dr. Moore for 



m 



he devoted to guidance and advice. 

The writer wishes to thank the curators of the following 
herbaria for the loan of specimens: ARIZ, B, bab, bh, bm, 

BR, C, CORD, DS, E, F, G, GH ? GOET, HAL, K, L, LA, M, MA, MICH, 
MO, MSC, NY, PH, R, RSA, P, SP, TENN, U, UC, US, VEN, VT, 

Wis. The staff at the Instituto de Biologia, Universidad 
Nacional Autonoma de Mexico (mexu), have been espe- 
cially kind and helpful during the author's trips to Mexico. 



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Rhodora [Vol. 77 



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Vegetation von El Salvador. Abh. Auslandsk., Reihe C, 
Naturwiss. 18: 98-179. 

. 1963a. Anatomische und histogenetische Untersuch- 

ungen an Laubsprossen und Bluten der Commelinaceen. Bot. 
Jahrb. Syst. 82: 1-99. 

• -. 1963b. 'One-sided' and 'two-sided' cincinni in the Com- 



melinaceae — a correction. Kew Bull. 17: 315-316. 

Rose, J. N. 1906. New names for two recently described genera 
of plants. Proc. Biol. Soc. Wash. 19: 96. 

Ruiz, H., & J. Pavon. 1794. Florae Peruvianae, et Chilensis Pro- 
dromus. 153 pp. Sancha, Madrid. 

Schlechtendal, D. F. L. 1831. Plantarum mexicanarum a eel. 
viris Schiede et Deppe collectarum recensio brevis auctoribus 
D. de Schlechtendal et Ad. de Chamisso. Linnaea 6: 22-64. 

. 1853. Plantae \Vag:enerianae Columbicae. Linnaea 26: 



127-144. 



Sinclair, C. 1968. Pollination, hybridization, and isolation factors 
in the erect Tradescantias. Bull. Torrey Bot. Club 95: 232-240. 

Standley, P. C, & J. A. Steyermark. 1944. Studies of Central 
American Plants. IV. Field Mus. Nat. Hist., Bot. Ser. 23: 31- 



109. 



& . 1952. Commelinaceae. In: Flora of 

Guatemala. Fieldiana: Bot. 24(3): 1-42. 

Stearn, W. T., & L. H. J. Williams. 1957. 'Martin's French 
Guiana plants and Rudg-e's "Plantarum Guianae rariorum 
icones." Bull Jard. Bot. fitat. 27: 243-265. 

Steyermark, J. A. 1963 ('1964') . Notes on Ecuador plants. Phy- 
tolotfia 9: 337-350. 

Tomlinson, P. B. 1966. Anatomical data in the classification of 
Commelinaceae. Jour. Linn. Soc, Bot. 53: 371-395. 

— . 1969. Anatomy of the Monocotyledons Vol. 3. Com- 
melinales-Zintfiberales. 446 pp. Clarendon Press, Oxford & 
London. 

Urban, I. 1906. Plantae novae andinae imprimis Weberbauerianae 

I. Bot. Jahrb. Syst. 37: 373-463. 
. 1912. Symbolae Antillanae. Vol. 7. 580 pp. 



1975] Tripogandra — Handlos 319 



Willdenow, C. L. 1799. Species Plantarum. 2(1): 1G-22. 
Woodson, R. E. 1942. Commentary on the North American genera 
of Commelinaceae. Ann. Missouri Bot. Gard. 29: 141-154. 

BOTANY DEPARTMENT 
UNIVERSITY OF ZAMBIA 
LUSAKA, ZAMBIA 



320 



Rhodora [Vol. 77 



Fig. 1. Upper pedicel and lower calyx of Tripogandra purpur- 
a8cens bearing capitate hairs. X 17. 

Fig. 2. Upper pedicel and lower calyx of T. mxicola bearing 
uniseriate hairs. X 17. 

Fig. 3. Upper pedicel and lower calyx of T. serrulata. X 17. 

Fig. 4. Sheathing leaf base and node of T. serrulata showing a 
continuous line of uniseriate hairs on the sheath and internode. 

X 11. 



1975] 



Tripogandra — Handlos 



321 







322 



Rhodora [Vol. 77 



Fig. 5. Outer stamen of Tripogandra pwrpuroAcens. X 11 

Fig, 6. Stamen of T. ample x arts. X 11. 

Fig. 7. Stamen of T. amplewcaulw. X 11. 

Fig. 8. Stamen of T. granciiflora. X 11. 

Fig. 9. Stamen of T. granciiflora. X 11. 

Fig. 10. Outer stamen of T. montana. X 11. 

Fig. 11. Stamen of T. guerrerensis. X 11. 

Fig. 12. Outer stamen of T. saxicola. X 22. 

Fig. 13. Stamen of T. palmeri. X 22. 

Pier. 14. Stamen of T. anguxti folia. X 22. 



1975] 



Tripogandra — Handlos 



*> o o 




324 



Rhodora t Vo1 - 77 






Fig. 15. Androecium of Tripogandra guerrerensis. X 11 

Fig. 16. Flower of T. serrulata. X 7.5. 

Fig. 17. Staminode of T. guerrerensis. X 11. 

Fig. 18. Inner stamen of 7\ purpuraxccns. X 11- 

Fig. 19. Inner stamen of T. serrulata. X 11. 

Fig. 20. Inner stamen of T. saxicola. X 11. 

Fisr. 21. Staminode of T. palmeri. X H. 

Fig. 22. Staminode of T. palmeri. X 11. 



1975] 



Tripogandra — Handles 



^9S 














326 Rhodora [Vol. 77 



Fig. 23. Distal portion of staminode of Tripogandra amplexicaulis 
showing the inflated filament. X 11. 

Fig. 24. Germinating seed of T. purpurnHcenn showing the emerg- 
ing radicle and capping embryotega. X 34.5. 

Fig. 25. Androecium and gynoecium of T. amplexans showing 
the relative positions of stamens and staminodes. X 11. 

Fig. 26. Staminode of T. amplexans, lateral view. X 11. 

Fig. 27. Distal portion of filament and anther of T. angustifolia. 
X 22. 

Fig. 28. Distal portion of staminode of T. grandi flora. X 11. 

Fig. 29. Staminode of T. amplezicatllis-T '. amplexans intermediate. 
X 11. 

Fig. 30. Staminode of T. angustifolia. X 16. 

Fig. 31. Distal portion of inner stamen of T. montana. X 11. 



1975] 



Tripotfandra — Handlos 



327 




328 






Rhodora 



[Vol. 



i i 



Fig. 32. Seed of Tripogandra multiflora, dorsal surface. 

Pig. 33. Seed of T. multiflora, ventral surface showing puncti- 
form hilum. 

Fig. 34. Seed of T. multiflora, dorsal surface. 

Fig. 35. Seed of T. multiflora, ventral surface showing puncti- 
form hilum. 

Fig. 36. Seed of Tradescantia triandra, dorsal surface. 

Fig. 37. Seed of T. triandra, ventral surface showing punctiform 
hilum. 

Fig. 38. Seed of Tripogandra serrulata, dorsal surface. 

Fig. 39. Seed of T. serrulata, ventral surface showing puncti- 
form hilum. 

Fig. 40. Seed of T. diuretica, dorsal surface. 

Fig. 41. Seed of T. diuretica, ventral surface showing puncti- 
form hilum. 

Fig, 42. Seed of T. glandulosa, dorsal surface. 

Fig. 43. Seed of T. glandulosa, ventral surface showing puncti- 
form hilum. 

Fig. 44. Seed of T\ montana, dorsal surface. 

Fig. 45. Seed of T. montana, ventral surface showing puncti- 
form hilum. 

Fig. 46. Seed of T. montana, lateral surface. 



Fig. 



47. Seed of T. brasiliensis, dorsal surface. 



Fig. 48. Seed of T. brasilierms, ventral surface showing elliptical 



hilum. 



Fig. 49. Cleared ovary of T. guerrerensis showing two seeds, the 
upper larger and the lower smaller, and the dorsal bundle of one 

carpel. 

The scale represents 1 mm. 



1975] 



Tripogandra — Handloj 



s 



329 













• 



i 



% 






- 



v 



% f - € ■ « 


















330 



Rhodora 



[Vol. 77 



Fig, 50. Seed of Tripogandra purpurascens subsp. purpurascens, 
dorsal surface. 

Fig. 51. Seed of T. purpurascens subsp. purpurascens, ventral 
surface showing: punctiform hilum. 

Fig. 52. Seed of T. disgrega, dorsal surface. 

Fig. 53. Seed of T. disgrega, ventral suface showing punctiform 
hilum. 

Fig. 54. Seed of T. purpurascens subsp. aus trails, dorsal surface. 

Fig. 55. Seed of T. purpurascens subs]), aus traits, ventral sur- 
face showing punctiform hilum. 

Fig. 56. Seed of T. saxicola, dorsal surface. 

Fig. 57. Seed of T. saxicola, ventral surface showing punctiform 
hilum. 

Fig. 58. Seed of T. amp lex arts, dorsal surface. 

Fig. 59. Seed of T. ample xans, dorsal surface. 

Fig. 60. Seed of T. amplexans, ventral surface showing linear 
hilum. 

Fig. 61. Seed of T. warming iana, dorsal surface. 

Fig. 62. Seed of T. amplexicaulis-T \ arnplexans intermediate, 
dorsal surface. 

Fig. 63. Seed of T* amplexicaulis-T. amplexctns intermediate, 
ventral surface. 

Fig. 64. Seed of T. amplexicaulis, dorsal side showing alveolate 

surface. 

Fig. 65. Seed of T. amplexicaulis, ventral side showing linear 
hilum and alveolate surface. 

Fig. G6. Seed of T. amplexicaulis, dorsal surface. 

Fig. 67. Seed of 7 1 . amplexicaulis, ventral surface showing linear 
hilum. 

The scale represents 1 mm. 



1975] 



Tripogandn 



t 



Handlos 



331 



*•■ 




























332 



Rhodora 



[Vol. 77 



Fig, 68. Seed of THpogandra gucrrcrensis, dorsal surface, lower 
seed of the locule. 

Fig. 69. Seed of T. guerrerensis, ventral surface showing ellipti- 
cal hilum, lower seed of the locule. 

Fig. 70. Seed of T. guerrerensis, dorsal surface, upper seed of 
the locule. 

Fig-. 71. Seed of T. guerrerensis, ventral surface showing ellipti- 
cal hilum, upper seed of the locule. 

Fig. 72. Seed of T. palmeri, dorsal surface. 

Fig. 73. Seed of T. palmeri, ventral surface showing- linear 
hilum. 

Fig. 74. Seed of T. angustifolia, dorsal surface. 

Fig- 75. Seed of T. avguxtifolia, ventral surface showing puncti- 
form hilum. 

Fig. 76. Seed of T. grandiflora, dorsal surface. 

Fig. 77. Seed of T. grandi flora, dorsal surface. 
Fig. 78. Seed of T. grandiflora, ventral surface showing linear 
hilum. 

The scale represents 1 mm. 



1975] 



Tripogandra 



Handlos 



333 








70 




76 













75 



i 1 . 




■ ■ 






THE FLORA OF CONCORD 1 : A REVIEW 



The greater Boston area has a long history of botanical 
publications, beginning with Bigelow's Florida Bostonicnsis 
of 1814. A "Flora of Middlesex County" by L. L. Dame 
and F. S. Collins was published in 1888. R. J. Eaton's Flora 
of Concord is in this tradition. Based on his own knowl- 
edge, the herbarium of the New England Botanical Club and 
the Gray Herbarium — which holds the most of Thoreau's 
herbarium — he has synthesized an annotated catalog which 
is both ecological and historical. 

The Flora of Concord has been a labor of love. It is for- 
tunate that the Museum of Comparative Zoology of Harvard 
University, having a Field Station in Concord, should have 
undertaken the publication of so useful a book. One hopes 
that this will stimulate the production of other local floras 
which are, after all, the stuff on which more synoptic works 
must eventually be based. 



GORDON P. DE WOLF, JR. 

ARNOLD ARBORETUM 

JAMAICA PLAIN, MASS. 021:50 



'Eaton, R. J. — A Flora of Concord; from Thoreau's Time to the 

Present Day. Cambridg-e, Mass.: Museum of Comparative Zoolopv: 
1974. 



334 



REPORT ON ENDANGERED AND THREATENED 
PLANT SPECIES OF THE UNITED STATES 



i 



The Smithsonian Institution's Report on Endangered and 
Threatened Plant Species of the United States' has been 
presented to Congress and published by the Government 
Printing- Office. Contained in the 200-page Report are lists 
of endangered, threatened, commercially exploited and re- 
cently extinct species of the United States (including Alaska 
and Hawaii), as well as recommendations for the preserva- 
tion and protection of these species. Listings of endangered 
and threatened species arranged alphabetically by States 
are included. A limited number of complimentary copies 
of the Report are available, upon written request, from: 
Endangered Flora Project, Department of Botany, Smith- 
sonian Institution, Washington, I). C. 20560. 



EDWARD S. AYENSU 
DEPARTMENT OF BOTANY 
SMITHSONIAN INSTITUTION 
WASHINGTON, D. C. 20560 



3 3 5 



APPEAL FOR SUPPORT FOR THE 
INDEX HOLMENSIS PROJECT 

The INDEX HOLMENSIS is an index of plant distribu- 
tion maps with world-wide coverage. It is the only interna- 
tional bibliography on distribution of vascular plants in 

area and vegetation maps. 

We have so far published four volumes, i.e. volume I, 
covering vascular cryptogams and gymnosperms, volume II 
containing Monocotyledoneae A-I, volume III Monocotyle- 
doneae J-Z, and finally volume IV covering Dicotyledoneae 
A-B, together more than 1000 pages. We intend to continue 
to publish one volume each year. The total number of 
distribution maps so far published is estimated at about 
400,000, all of which will finally be listed in the index 
or its supplement. Although the main work is at present 
done at the Swedish Museum of Natural History in Stock- 
holm, the indexing work is served by an international 
editorial board. Members of this board to some extent 
guarantee completeness of the file for their particular area. 

Still, the number of area and vegetation maps published 
annually is growing rapidly owing to the increased im- 
portance that is felt for the geographic compound of plant 
taxa. Consequently, not only are there wide areas all over 



system 



maDDiner has become 



in different fields, as for instance economic botany, paleo- 
botany, vegetational history, palynology, and last but not 
least phytocoenology. 



In order 



HOLMENSIS 



and its planned supplement volumes up-to-date we here- 
with ask our fellow botanists to inform us about their 
published maps and/or to send reprints of their publica- 
tions. Needless to say, we shall also continue to supply all 



maps 



HOLMENSIS 



All correspondence should be addressed to: Hans Tralau, 
The Swedish Museum of Natural History, S — 104 05 
Stockholm 50. 



Volume 77, No. 810, including pages 165-3S6, was issued Sept. SO, 1975 



336 



Cover illustration 

Rhododendron canadense (L.) Torr. 

from 
Curtis' Botanical Magazine t.474 



RHODORA June, 1975 Vol. 77, No. 810 



CONTENTS 



De Plantis Toxicariis E Mundo Novo Tropicale Commentationes 
XII. Notes on Biodynamic Piperaceous Plants 

Richard Evans Sckultes 165 



Chromosome Counts of Compositae from the United States, 
Mexico, and Guatemala 

David J. Keil and Tod F. Stuessy 

A Vegetational Survey of the Vascular Plants of the Kent 
Island Group, Grand Manan, New Brunswick 
John W. McCain 

A Red-petioled Form of Thalia geniculata L. from Central 

Florida 

Allen G. Shuey .... 

The Taxonomy of Tripogandra (Commelinaceae) 
Wayne L. Handlos 

The Flora of Concord: A Review 

Gordon P. DeWolf, Jr 

Report on Endangered and Threatened Plant Species of the 
United States 

Edward S. Ay&nsu 

Appeal for Support for the Index Holmensis Project 

Hans Tralau - 



171 



196 



210 



213 



334 



335 



336 




JOURNAL OP THE 



NEW ENGLAND BOTANICAL CLUB 




Vol. 77 



September, 1975 



No. 811 












Stlfe ^Nefo ^wgkmb ^tfiamcal Club, ^ttc 

Botanical Museum, Oxford Street, Cambridge, Mass. 02138 



Conducted and published for the Club, by 
ALFRED LINN BOGLE, Editor-in-Chief 

ROLLA MILTON TRYON 
STEPHEN ALAN SPONGBERG 
GERALD JOSEPH GASTONY 
RICHARD EDWIN WEAVER 



Associate Ed i tort 



RHODORA. — A quarterly journal of botany, devoted primarily to 
the flora of North America and floristically related areas. Price 
$20.00 per year, net, postpaid, in funds payable at par in the United 
States currency at Boston. Some back volumes, and single copies 
are available. For information and prices write RHODORA at 



address given below. 
Scientific papers a 



be 



mittee for publication. Articles concerned with systematic botany 
and cytotaxonomy in their broader implications are equally accept- 
able. All manuscripts should be submitted in duplicate, and should 
be double-spaced throughout. Please conform to the style of recent 
issues of the journal. Illustrations can be used only if the cost of 
engraver's blocks is met through the author or his institution. 
Forms may be closed five weeks in advance of publication. Ex- 
tracted reprints, if ordered in advance, will be furnished at cost. 



proofs 



Bogl 



Dept. of Botany and Plant Pathology, 
University of New Hampshire, 
Durham, New Hampshire 03824 

Subscriptions and orders for back issues (making all remittances 
payable to RHODORA) should be sent to RHODORA, Botanical 
Museum, Oxford Street, Cambridge, Mass. 02138. In order to receive 
the next number of RHODORA, changes of address must be received 
prior to the first day of March, June, September or December, 

Second Class Postage Paid at Boston, Mass. 



MANUFACTURED BY 
THE LEXINGTON PRESS, INC. 
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IRbbfcora 



JOURNAL OF THE 

NEW ENGLAND BOTANICAL CLUB 



Vol. 77 September, 1975 No. 811 



A REVISION OF THE 
NORTH AMERICAN GENUS AMORPHA 

(LEGUMINOSAE-PSORALEAE) 



Robert L. Wilbur 



l 



Amorpha is a small North American genus of papilionate 
legumes belonging to the tribe Psoraleae. It has been the 
subject of several taxonomic revisions (Schneider, 1907; 
Rydberg, 1919; and Palmer, 1931) as well as numerous 
regional treatments and one might suppose that such atten- 
tion would have resulted in a rather satisfactory taxonomic 
understanding at the present time. However, such a favor- 
able supposition would be completely unjustified and my 
presentation below is offered as an alternative. Although 
it is the result of an intensive if somewhat sporadic study 
timewise for more than fifteen years and an examination 
of more than twelve thousand specimens, I am not so naive 
as to believe that my account of the genus presented below 
is the ultimate answer either. Naturally I hope that future 
studies will show that the revision presented here is a 
closer approximation of the biological reality than the 
earlier accounts, but I would be less than candid if I were 
not to admit that I have found the genus to be extremely 
difficult and not especially tractable to the usual investiga- 



tory 



experimental" taxonomi 



'Grateful acknowledgment is hereby made for support given by the 
National Science Foundation to this research under NSF Grants 

5636 and GB-138ir). 



337 






38 Rhodora [Vol. 77 



The difficulty encountered with Amorpha as with any 
taxonomically perplexing taxon is not that there is too 
much variability or too little but that the variability that 
seemed so discrete when a relatively small number of speci- 
mens was examined becomes ever so much more like a 
continuum when thousands of specimens are studied. En- 
vironmentally induced plasticity of at least certain members 
of the genus is sometimes spectacular, and it certainly 
would prove disconcerting to those who have recognized 
numerous taxa in what I refer to as the frvticosa-complex 
to see the number of specimens that clearly possess the 
characteristics of one "taxon" in the growth produced 
early in the season and that of another "species" or "va- 
riety" in the later growth. The extraordinary morphologi- 
cal diversity exhibited by certain species of Amorpha, due 
both to apparently great genetical flexibility (i.e. numerous 
biotypes) and environmental plasticity, has resulted in the 
recognition of numerous taxa. For example, Rydberg 
(1919) recognized 23 species while Palmer (1931) ac- 
cepted 20 species, 11 additional varieties and 7 forms for 
a total of 38 named taxa. The treatment presented here 
recognizes 15 species (including one new species, A. oua- 
chitensis Wilbur) and 3 additional varieties. 

Any definitive statement concerning the interrelation- 
ships of the genus Amorpha with its tribal neighbors in 
the Psoraleae must await more complete understanding of 
those genera as well as of Amorpha itself. Morphologically 
Amorpha seemingly is most closely related to the ditypic 
Parryella T. & G. ex A. Gray of Arizona and New Mexico 
and to the more widespread but still western genus Eysen- 
hardtia H.B.K., which ranges from central Texas west to 
California and south into Guatemala. Parryella differs 
principally from Amorpha in that it is without at least all 
outward evidence of petals while Eysenhardtia has all 5 
petals. The fruits of all three genera are 1(2) -seeded but, 
although all are indehiscent, those of Parryella and Amor- 
pha are far more similar in texture, shape, and all other 
respects. With the facts available to us at the present time, 



1975] Amorpha — Wilbur 339 

future research may support the hypothesis that Amorpha, 
Parry ella, and Eysenhardtia were derived from a common 
ancestral complex and that Amorpha and Parry ella are 
more closely related to one another than either is to Eysen- 
hardtia. 

My gratitude to the curators of the herbaria listed below 
is far greater than is even usually the case in similar ac- 
knowledgments for they have been extremely tolerant in 
permitting me to keep loans for far longer than is custom- 
ary and without their understanding the study could not 
have been completed. 

A, ARIZ, CAS, COLO, CS, DES, DS, DUKE, F, FLAS, FSU, GA, 
GH, IA, ISC, JEPS, KANU, KSC, MICH, MINN, MO, NCU, NDA, 
ND-GREENE, NEB, NSC, NY, OKL, OKLA, PHIL, POM, RSA, SDC, 
SDU, SMU, TENN, TEX, UARK, UC, UMO, US, USF. 



Amorpha L., Sp. PI. 743. 1753; Gen. PI. ed. 5. 319. 1754. 

Bonfidia Necker, Elem. Bot. 3: 46. 1790. (Art. 20(2) 
of the I.C.B.N. clearly states that this is not to be 
regarded as a generic name.) 

Suffrutescent low shrubs to large bushy-topped shrubs. 
Leaves odd-pinnately compound with .setaceous to linear, 
caducous stipules and short- to long-petiolate ; leaflets (7) 
11-29(45), entire to crenulate, epunctate to very densely 
and conspicuously glandular punctate, petiolulate, stipu- 
late on the upper side. Inflorescence a spike-like, terminal 
raceme or the racemes clustered and then appearing panicu- 
late; flowers pedicellate from the axil of a setaceous to 
linear, caducous bract. Calyx persistent, obconic, funnel- 
form or even shortly campanulate, almost imperceptibly to 
strongly 5-lobed. Corolla reduced to a single petal, the 
banner, and it erect, enveloping the internal floral series, 
clawed, obovate to obcordate, often eventually the apical 
lobes adaxially recurved, purple, blue, violet or white, entire 
to erose. Stamens 10, at least basally monadelphous and 
occasionally the staminal tube exserted beyond the calyx; 
the filaments distinct above and exserted beyond the calyx 



340 Rhodora [Vol. 77 



and usually the petal; anthers versatile, small, oblong, 



longitudinally dehiscent. 



compressed 



beyond 



and often the petal; stigma terminal, capitate. Fruit a 
1-seeded, indehiscent pod, ± oblique, straight to strongly 
curved, compressed, slightly to very noticeably longer than 
the calyx, eglandular to conspicuously punctate glandular; 
seeds ovoid to oblong, laterally compressed. 

Typification : Amorpha fruticosa L. ; the genus was com- 
prised of only one species when originally established. 

Distribution : a North American genus of about 15 spe- 
cies ranging from southern Canada south to southern 
Florida and west to California. 






KEY TO THE SPECIES OF AMORPHA 

1. Petioles and the rachises of leaves beset with spine-like 

glands; plants of Arizona, California and Baja Cali- 
fornia. 

2. Vexillum reddish-purple, claw indistinct but about 
1 mm long; filaments 6-7 mm long, united for the 
basal 2-3.2 mm; fruit densely to moderately pilosu- 
lose throughout (except in one area north of San 
Francisco Bay) ; plants of Arizona and northern 
California south into northern Baja California. . . 



1. A. calif 



2. Vexillum white, claw 2-3 mm long; filaments 8-12 
mm long, united for the basal 1-1.5 mm; fruit gla- 
brous or sparingly pilosulose only near the apex; 
plants known only from the Sierra San Pedro Martir 
of northern Baja California 2. A. apiculata. 

1. Petioles and the rachises of leaves lacking spine-like 
glands. 

t 

3. Shrubs usually less than 1 m tall; leaves shortly 
petiolate with the petiole typically shorter than the 
width of the lowest leaflet. 



1975] Amorpha — Wilbur 341 



4. Midvein of the leaflets included or but barely 
exserted and then its tip distinctly swollen, or if 
noticeably exserted, then the tip of mucro con- 
spicuously swollen ; plants of the southeastern 
United States. 

5. Calyx-tube densely puberulent, or, if not, at 
least the lobes externally sparsely puberulent; 
stem, rachises (of both leaf and inflorescence), 
and leaflets densely pubescent, or, if not, usu- 
ally at least somewhat puberulent; leaflets 
entire or but inconspicuously crenulate; peti- 
olules usually less than 1.5 mm long* and 
densely to sparsely pubescent; ranging from 
North Carolina south into peninsular Flor- 



ida 



3. A. hcrbacea. 



5. Calyx-tube and usually the lobes glabrous; 
stem, rachises (of both leaf and inflorescence), 
and leaflets glabrous or but very sparsely 
puberulent or short-pubescent ; leaflets usually 
conspicuously crenulate; petiolules usually 
more than 1.5 mm long and glabrous (or 
rarely very sparingly puberulent) ; known 

only from Dade County, Florida 

4. A. crenulata. 



4. Midvein of the leaflets exserted into a slender 
mucro with a tapered tip; plants of the south- 
eastern United States and much of the central 
part of the continent. 

6. Adaxial calyx-lobes 0.1-0.8(1.8) mm long, 
the abaxial lobe (1)1.2-1.6(1.8) mm long; 
plants of the coastal plain of the southeastern 
United States (the Carolinas and Georgia). 

5. A. georgiana. 

6. Adaxial calyx-lobes (0.6)1-1.5(1.8) mm long, 
the abaxial lobe (1.2)1.5-2.2(2.5) mm long; 
plants of the central United States (southern 
Canada south into Texas and from Indiana 
west into New Mexico and Wyoming). 



342 Rhodora [Vol. 77 



7. Foliage and/or calyces conspicuously pu- 
bescent to the unaided eye and often ca- 
nescent 6. A. canescens. 

7. Foliage and calyces glabrous or nearly so, 
or at least not conspicuously pubescent to 
the unaided eye and never canescent. 

8. Leaflets appearing epunctate or at 
least the punctate glands on the lower 
surface of the leaflets not discernible 
without magnification ; racemes usually 
clustered and mostly in groups of 5-10. 
6. A. canescens. 

8. Leaflets conspicuously punctate and 
the glands readily visible without mag- 
nification ; racemes solitary 



7. A. nana. 

3. Shrubs usually more than (1)1.2 m tall; leaves with 
petioles longer than the width of the lowermost 
leaflet 

9. Calyx-lobes nearly obsolete, all less than 0.8 mm 
long and never narrowly acute. . . 8. A. glabra. 

9. Calyx-lobes or at least some well developed and 
longer than 0.8 mm and some narrowly acute or 
acuminate. 

10. Calyx-lobes from about half as long to as 

long as the calyx-tube, all acute or acumi- 
nate; the lateral lobes more than 1 mm long. 
11. Secondary venation slightly to moder- 
ately elevated beneath but the leaflets 
never conspicuously reticulate; abaxial 
calyx-lobes (1.8)2.5-3.5 mm long; plants 
of the Carolinas, Georgia, and Alabama. 

9. A. schwcrinii. 

11. Secondary venation conspicuously ele- 
vated beneath often to the 3rd and 4th 
degree of branching and thus forming a 
striking reticulum; abaxial calyx-lobes 



1975] Amorpha — Wilbur 343 



(1.2)1.5-2.0 mm long; plants of Louisi- 



ana and eastern Texas 



10. A. paniculata. 

10. Calyx-lobes or at least some of them less 

than half as long as the calyx-tube, at least 
the adaxial usually rounded; the lateral 
lobes less than 1 mm long. 
12. Foliage, calyces and pedicels blackening 

or at least darkening upon drying; 
calyx-tube eglandular to sparingly 
glandular with very small and incon- 
spicuous, slightly elevated punctate 

glands 11. A. nitens. 

12. Foliage, calyces and pedicels never 

blackening and rarely even darkening 
upon drying; calyx-tube usually glandu- 
lar with often large, conspicuous, ele- 
vated punctate glands. 
13. Petiolules conspicuously pustulate- 
glandular ; leaflets typically emargi- 
nate and often with a globose tip 
to the midvein; plants of Texas or 
Oklahoma and/or Arkansas. 
14. Petals present on specimen. 

15. Petals bright blue to deep 

violet; stipules glabrous; 
plants of eastern Texas 



and Oklahoma 



12. A. laevigata. 

15. Petals purplish; plants of 

central Texas ; stipules 
pubescent ; eastern Okla- 
homa and/or west central 

Arkansas. 

16. Abaxial calyx-lobes 

(0.8)1.0-1.2(1.4) mm 

long ; central Texas 

on the Edwards Pla- 



344 Rhodora [Vol. 77 



teau. 



. . 13. A. roemeriana. 
16. Abaxial calyx-lobes 

about 0.6-0.9(1.2) 
mm long; Ouachita 
Mts. of southeastern 
Oklahoma and west 
central Arkansas. . . 
. 14. A. otmchitensis. 
14. Petals lacking on specimen. 

17. Calyx-tube beset with 

large conspicuous pustu- 
late glands throughout 
the upper 1/2-2/3 its 
length, glabrous; eastern 
Texas and southeastern 
Oklahoma 

12. A. laevigata. 

17. Calyx-tube sparingly to 

moderately supplied with 

often inconspicuous pus- 
tulate glands in the up- 
per third of its length; 
glabrous to moderately 
spreading to appressed 
short-pubescent ; central 
Texas or the Ouachita 
Mts. of southeastern Ok- 
lahoma and west central 
Arkansas. 

18. Abaxial calyx-lobes 

(0.8)1.0-1.2(1.4) mm 
long; fruit about 6-7 
mm long and 2.5-3.5 
mm wide ; central 
Texas on the Ed- 
wards Plateau 

. 13. A. roemeriana. 



1975] Amorpha — Wilbur 345 



18. Abaxial calyx-lobes 

0.6-0.9(1.2) mm long; 
fruit about 7-9 mm 
long and 3-4 mm 
wide ; Ouachita Mts. 
of southeastern Okla- 
homa and west cen- 
tral Arkansas 

. 14. A. ouachitensis. 
13. Petiolules eglandular or with only 

inconspicuous glands; leaflets acute 
to broadly rounded, only rarely 
emarginate and the midvein only 
rarely terminating in a globose or 
swollen tip; plants of almost the 

entire range of the genus 

15. A. fruticosa. 



1. Amorpha californica Nutt. in Torr. & Gray, Fl. N. Am. 1: 

306. 1838. 

Erect, bushy shrub (0.7)1-3(4) m tall. Current season's 
growth sparingly to more typically moderately puberulent 
or crinkly-pilosulose bearing few to numerous, usually con- 
spicuous, often apically pointed or sharply hooked, as well 
as basally swollen, resiniferous, amber-colored glands. 
Leaves ascendant to spreading, (0.5)1-1.5(2.0) dm long. 
Petioles (0.7)1-1.5(2.5) cm long, usually equaling or longer 
than the width of the lowermost leaflet, densely crinkly- 
pilosulose or puberulent to glabrous with several to numer- 
ous, rounded and pointed, resiniferous, pustulate glands. 
Stipules caducous, oblong-lanceolate to lanceolate or linear, 
eglandular, reddish-brown, sparingly to densely, appressed 
tawny-pilosulose, (2)4-6 mm long. Rachis of leaf slender, 
about 1 mm in diameter, densely crinkly-pilosulose or 
puberulent to glabrous, moderately beset with both sharp- 
pointed and rounded, amber-colored, resiniferous, sessile 
glands. Leaflets (11)13-19(25), oblong to elliptic-oblong, 
(0.8)1.5-2.5(4.2) cm long, (0.6)1-1.5(2.0) cm wide, typi- 



346 Rhodora [Vol. 77 



cally 1.5-2.2 times as long as wide, opposite or more typi- 
cally alternate, symmetrical, more or less broadly rounded 
to subcordate basally, typically broadly rounded to obtuse 
and emarginate apically, entire to inconspicuously crenu- 
late; secondary veins very slightly, if at all, elevated be- 
neath. Midvein usually terminating in a swollen, globose 
tip flush with the margin or very rarely slightly exserted 
or very rarely somewhat tapering. Lower surface of leaf- 
lets moderately pilosulose throughout but densely so along 
the midvein to glabrous or very nearly so, conspicuously 
glandular-punctate below with light to dark brown, resin- 
ous glands of apparently 2 sizes; upper surface typically 
moderately pilosulose but occasionally glabrous, eglandu- 
lar. Petiolule 0.7-1.5(1.8) mm long, densely spreading 
pilosulose to glabrous, sparingly to moderately pustulate- 
glandular. Racemes erect, usually solitary but rarely few 
together, (0.5)1-1.8(2.5) dm long; rachis of inflorescence 
moderately spreading-pilosulose, hirsutulous or puberulent 
to glabrous, sparingly glandular pustulate with both 
rounded and sharp-pointed glands. Pedicels 0.3-1.2 mm 
long, densely spreading- to appressed-pilosulose hirsutulous 
or puberulent to glabrous, eglandular; bracts lanceolate to 
narrowly linear, 1.5-4 mm long, externally moderately 
spreading- to appressed-pilosulose, eglandular to 
ately pustulate-glandular, internally glabrous, caducous. 
Calyx-tube narrowly cylindric or somewhat funnelform, 
2.0-2.8 mm long, moderately spreading- to appressed-pilosu- 
lose or puberulent throughout to glabrous, abundantly to 
sparingly pustulate-glandular in upper half. Calyx-lobes 
usually tipped by a conspicuous pustulate gland and also 
frequently sparingly to moderately glandular-pustulate, 
densely to moderately pilosulose or glabrous both externally 
and internally. Adaxial calyx-lobes broadly to narrowly 
triangular-dentate, acute, (0.4)0.8-1.5(2) mm long; lateral 
calyx-lobes slightly longer, acute to acuminate, (0.5)1-2 
(2.2) mm long; abaxial lobe narrowly lanceolate, acumi- 



mo 



nate, (0.8)1.5-2.5 mm 



mm 



mm 



1975] Amorpha — Wilbur 347 



short, indistinct claw about 1 mm long, moderately to 
strongly arching, apically more or less truncate to emargi- 
nate and often with a central apiculation, reddish-purple 
and with the claw and blade both strongly enveloping the 
filaments. Filaments 6-7 mm long, united into a sheath for 
the basal 2.0-3.2 mm, glabrous; anthers about 0.5-0.7 mm 
long, pale yellow. Style moderately ascending-pilose; ovary 
either densely pilosulose or very sparingly so and then only 
apically. Fruit 6-8 mm long, 2.5-3.5 mm wide, broadest 
above the middle, tapering somewhat basally, with the 
adaxial margin straight or slightly outwardly outcurved 
above and the abaxial strongly outwardly bowed, densely 
to moderately pilosulose throughout or only sparingly 
strigillose apically, pustulate-glandular in the upper half 
or two-thirds. Seeds smooth, plump, 2.5-3.2 mm long, light- 
to olive-brown. 

Distribution: Northern California (approximately -11° 
N) south into the Sierra San Pedro Martir of northern 
Baja California (approximately 31° N) and sporadically 
eastward as far as southeastern Arizona. 

Two rather strikingly distinct varieties have been recog- 
nized within this species. Their distribution as shown by 
our rather conventional but crude mapping 1 does not sug- 
gest that they are strictly allopatric but a more refined 
appraisal of their relationship should wait at least careful 
field observations. The two may be separated by the fol- 
lowing key. 

A) Upper surface of leaflets, rachises and petiolules moder- 
ately to densely spreading-pilosulose or puberulent; 
calyx-tube moderately spreading-pilosulose or occasion- 
ally appressed-pilosulose ; calyx-lobes all, or at least the 
longest, over 1 mm long la. var. calif arnica. 

A) Upper surface of leaflets, rachises and petiolules gla- 
brous or very nearly so; pubescence on lower surface 
of leaflet lacking or largely restricted to the midvein ; 
calyx-tube glabrous or nearly so or sparingly to rarely 



348 Rhodora [ Vo1 - 77 



moderately strigillose ; calyx-lobes all 1 mm long or less. 



la. A. californica var. californica 



lb. var. napcnsis 



f 



1: 306. 1838. typification : "St. Barbara, Califor- 



Nuttall 



A. hispidula Greene, Fl. Fran. 14. 1891. typifica- 
tion: California: Monterey co., Hickman s.n. 



(lectotype, nd-g!). 
californica var. hispidula 
Arnold Arb. 12: 163. 1931. 



(Greene) Palme 



more than 1 mm 



more 



Branchlets, rachises, petioles and petiolules moderately 
spreading pilosulose or puberulent; leaflets spreading- to 
appressed-pilosulose above and below. Calyx-tube usually 
moderately to densely spreading- or appressed-pilosulose or 
puberulent, eglandular or sparingly and inconspicuously 
pustulate-glandular. Calyx-lobes \ 
long. Pod moderately to densely 
short-pubescent. 

Distribution : Coastal Ranges north of San Francisco to 
about 38°30' N and along the Coastal Ranges south of San 
Francisco from Monterey Co. (about 36°30' N) into north- 
ern Baja California in the Sierra San Pedro Martir (about 
30°30' N) ; also in an area south of Mt. Shasta (about 
41° N) and on the western slopes of the Sierra Nevada 
(about 39° N) ; disjunctly occurring in central and south- 
eastern Arizona. (Map 2.) 



lb. 



A. californica var. napensis Jepson, Man. Fl. PI. Calif. 
556. 1925. TYPIFICATION: California: Howell Mt., 
Napa Range, Jepson 6835 (lectotype, jeps!). The 
lectotype, designated by Jepson (Fl. Calif. 2: 331. 
1936), is a sterile specimen and apparently a crown 
sprout while his number 6834 collected on the same 
day and the same place, is the fruiting specimen from 
which most of the diagnostic information was ob- 
tained. 



1975] 



Amorpha — Wilbur 



349 





MAPS 1-2. Map 1. Amorpha calif ornica var. napennix (dots 
north of San Francisco Ray) and A. apiculata (dots with horizontal 
line in Raja California). Map 2. A. calif ornica var. calif ornica. 



Branchlets, rachises, petioles and petiolules very spar- 
ingly pilosulose or more typically glabrous; leaflets gla- 
brous above and either glabrous beneath or the pubescence 
primarily restricted to the midvein. Calyx-tube glabrous 
or sparingly strigillose above or rarely moderately strigil- 
lose throughout. Calyx-lobes usually all 1 mm long or less 
and ranging from 0.4-1.0 mm long. Fruit sparingly to 
moderately strigillose apically or throughout. 

Distribution : endemic to the area north of San Francisco 



Bay in M 
part.) 



Napa and Sonoma Counties. 



M 



1, in 



Both recent California floras (Munz 



1944) indicate the species is divisible into two varieties: a 

the Shasta area of southern 



northern one 



om 



Monterey 



ing from the Santa Lucia Mountains just south of Mon- 
terey south to the Santa Ana and Santa Rosa Mountains 



350 



Rhodora [Vol. 77 



of extreme Southern California. The range of the varieties 
presented by these authors is in obvious conflict with those 
found in this study although the morphological features 
characterizing the taxa in all three studies seem to be 
identical. 



2. Amorpha apiculata Wiggins, Contr. Dudley Herb. 1 : 171. 

1933. typification : Mexico: banks of the Rio Santo 
Domingo about 5 miles above Mission Santo Domingo, 
Baja California, Wiggins & Demaree 4776 (holotype, 
DS, not seen; isotypes, ARiz!, ds!; f!; ny!, pom!, rsa!). 

Erect, slender shrub 2-5 m tall. Current season's growth 
moderately to sparsely spreading-pilosulose bearing few to 
numerous, rounded, resiniferous, amber-colored, elliptic 
glands and a few, apically pointed, basally rounded, spine- 
like glands. Leaves ascendant to spreading, 1-2 dm long. 
Petioles 1-2.5 cm long, usually equaling or longer than the 
width of the lowermost leaflet, sparingly to moderately 
spreading-pilosulose, with several to numerous, usually 
rounded but occasionally pointed, amber-colored glands. 
Stipules caducous to tardily deciduous, linear to linear- 
setaceous, glandular, sparingly pilosulose, 2-3 mm long. 
Rachis of leaf slender, about 1 mm in diameter, sparingly 
to moderately pilosulose, moderately beset with rounded or 
pointed, amber-colored, resiniferous, sessile glands. Leaf- 
lets (9)13-19(25), elliptic to oblong-elliptic, mostly 1.5- 
3.0(3.8) cm long and (0.5)1.0-1.5 cm wide, typically 2.2- 
3.3 times as long as wide, usually alternate and symmetri- 
cal, tapering either both basally and apically and often 
acute to more or less rounded ; secondary veins very slightly 
elevated beneath. Midvein exserted, 0.2-0.8 mm long, 
slightly to conspicuously swollen at the tip. Lower surface 
sparsely pilosulose throughout or more or less restricted to 
the midvein where tending to be wide-spreading, "light 
green, slightly glaucous," conspicuously glandular-punctate 
below with dark, resinous glands of apparently 2 sizes; 
upper surface glabrous, "bright green," moderately to 
sparingly punctate-glandular. Petiolules 1.5-2.2 mm long, 



1975] Amorpha — Wilbur 351 



sparingly to moderately spreading to somewhat appressed 
pilosulose, moderately pustulate-glandular. Racemes soli- 
tary to several, 1-7 in number, 1-3 dm long, rachis mod- 
erately spreading-pilosulose; pedicels 0.8-1.2 mm long, 
moderately spreading to appressed pilosulose; bracts nar- 
rowly linear, 2.5-3.0 mm long, sparingly pilosulose, pustu- 
late-glandular, caducous. Calyx-tube narrowly funnelform, 
2.5-3.0 mm long, moderately spreading-pilosulose through- 
out, conspicuously pustulate-glandular in upper fifth. Calyx- 
lobes usually tipped by a resinous, pustulate gland and 
otherwise eglandular, moderately to sparingly pilosulose 
or puberulent both externally and internally. Adaxial caly- 
cine lobe triangular-dentate, acute to less frequently acumi- 
nate, 0.5-1 mm long; lateral lobes longer and more nar- 
rowly lanceolate, acuminate, 0.8-1 mm long; abaxial lobe 
narrowly lanceolate, acuminate, 1.2-2 mm long. Vexillum 
5-7 mm long, about 4 mm wide, broadly obovate with a 
slender claw and a broad, truncate apical notch with a 
centra], triangular-dentate apiculation about 0.1-0.2 mm 
long, white; the claw enveloping the filaments and the 
blade either plane or reflexed away from the filaments. 
Filaments greatly elongate, 8-12 mm long, united for the 
basal 1-1.5 mm, glabrous; anthers about 0.5 
parently pale yellow. Style moderately ascending-pilose; 
ovary glabrous except for the uppermost pilosulose region 
near the style. Fruit (apparently still immature) about 
(i mm long and 2.5 mm wide, broadest above the middle, 
tapering to base, with the adaxial margin very nearly 
straight and the abaxial margin gradually outwardly 
bowed, glabrous or sparingly pilosulose only near the top, 
conspicuously glandular-pustulate. 

Distribution: Known only from the Sierra San Pedro 
Martir of northern Baja California, Mexico at approxi- 
mately 31° N. (Map 1, in part.) 

This species is clearly very closely related to Amorplia 
californica and especially to that species's var. califoi-nica 
which apparently also reaches its southern limit in the 



mm 



352 



Rhodora [Vol. 



1 1 



Sierra San Pedro Martir. I have examined only eleven 
different collections of either species from this area. These 
two species have been confused both in the field and the 
herbaria by collectors and identifiers and at the present 



ot form a: 
angement 



seem 



om 



lit 



higher elevations. 



seemin 



om 



Additional collections and field observations from this re- 



much 



seems in some respects 



strikingly different from A. calif ornica var. calif ornica 
than is A. calif ornica var. napensis, nothing would seem to 
be gained by attempting to surmise now the results of the 
needed research. The three taxa form a most distinctive 
complex within the genus. 

3. Amorpha herbacea Walt., Fl. Car. 179. 1788. 



mos 



m hierh arising: from 



thick, woody, often horizontal rootstock reportedly up to 
2 m in length and 2 cm in diameter. Current season's 
growth dull reddish-purple, olivaceous to light brown, 
slender, about (1)2-4 mm in diameter, with numerous fine 
longitudinal grooves and ridges and sprinkled throughout 
with few to numerous, usually inconspicuous, small punc- 
tate glands, mostly densely short-pubescent or puberulent 
and then often appearing canescent but occasionally (espe- 
cially in Florida) only very sparingly pubescent. Leaves 
numerous, spreading to ascendant, mostly about (0.6)0.8- 
1.8(2.4) dm long. Petioles mostly densely short-pubescent, 
puberulent or strigillose but occasionally glabrous or nearly 
so and with few to numerous postulate glands especially 
abundant near the base, typically shorter than the width 
of the lowest leaflet but occasionally as long as 1.5 times 
the width, about (0.5)1-10(13) mm long. Stipules incon- 



1975] Amorpha — Wilbur *o* 

spicuous, pigmented like the stem and rachis, glabrous to 
■sparingly pubescent, caducous or persisting as a withered 
remnant, appressed, acicular to setaceous, about (1)1.2- 
2 5(3) mm long. Rachis of leaf slender, about 0.5-1 mm in 
ui««ic*cx- usually densely short-pubescent or puberulent 
throughout (or if differentially so, then more densely 
pubescent above in and about the channeled groove) to less 
commonly sparsely short-pilose, strigillose or even glabrous 
or nearly so, sparingly glandular throughout. Leaflets (lo) 
23-45(63) in number, about (0.7)1.0-2.5(3.2) cm long and 



imete 



(3)4-10(15) mm wide, mostly (1.2)2-3(4) times as long as 
broad, usually symmetrical but rarely asymmetrical, oppo- 
site or alternate, the interval between petiolules on the 
same side of the rachis about (0.3)0.6-1.2(1.8) cm long. 
Blades of leaflets mostly broadly to narrowly oblong or 
occasionally oblong-elliptic to elliptic, or rarely ovate-ob- 
long, ovate, obovate, obcordate or almost orbicular with the 
base usually broadly rounded to obtuse, occasionally sub- 
cordate, truncate or nearly so, oblique or rarely tapering 
and then more or less acute; the apex obtuse to broadly 
rounded to almost truncate and occasionally shallowly 
emarginate, with the midvein terminating in a sessile or 
subsessile, distinctly swollen, often globose, glandular knob, 
or when exserted, the swollen tip on a stalk about 0.Z- 
5(0 8) mm long. Texture of leaflets coriaceous to sub- 
coriaceous when dry, finely reticulate above from the very 
slightly elevated venation excepting the scarcely depressed 
midvein; the margin usually slightly to conspicuously invo- 
lute and very inconspicuously crenulate to entire, or nearly 
so Midvein prominently elevated beneath and the second- 
ary veins but slightly raised. Both surfaces of the leaflets 
usually very densely pubescent, but rarely the upper or 
even both sides glabrous or very nearly so, and when most 
nearly glabrous, then often strigillose beneath along the 
principal veins; pubescence of the upper surface usually 
short-pilose, puberulent, or even strigillose and that of the 
lower surface usually short-villous, pilose, puberulent, or 
rarely strigillose; punctate glands on the lower surface 



354 Rhodora [Vol. 77 



usually conspicuous and readily apparent to the unaided 
eye, appearing- to be of approximately two size classes, 
usually numerous but occasionally few, ranging in number 
from about (40)75-225(425) glands per half-leaflet. Petio- 
lule about (0.7)1-2 mm long, usually densely puberulent, 
occasionally strigillose, less commonly sparsely puberulent 
or strigillose and rarely glabrous, with few to numerous 
pustulate glands. Stipels acicular to setaceous, mostly (0.4) 
0.8-2 mm long, rarely tardily deciduous, glabrous to pu- 
berulent, especially near the base. Racemes usually few to 
several or occasionally numerous, appearing terminally 
clustered or occasionally solitary, about (1)4-12(20) in 
number, mostly appearing slender and elongate, about 
(0.3)1-1.8(4) dm long and (0.8)1.2-1.8(2) cm in 
sessile, subsessile, or on a peduncle 1-4(7) cm long and 
with a usually densely but occasionally sparsely puberulent, 
conspicuously ridged and grooved, sparingly pustulate 
glandular rachis bearing numerous, usually densely clus- 



diameter 



m 



casionally glabrous pedicels about (0.2)0.4-1.2(1.8) mm 
long, subtended by a caducous, sparsely glandular-punctate, 
usually sparingly puberulent to short-pubescent, brownish 
to amber colored, linear-subulate to setaceous bract about 
(1.2)1.8-2.5(3) mm long. Calyx-tube turbinate to nar- 
rowly campanulate or cylindrical, mostly about 1.5-2.5(3) 
mm high with the abaxial side slightly longer than the 
adaxial, usually either very densely puberulent or short- 
pilose throughout, and with the pubescence curly and 
spreading or, rarely, densely to sparingly minutely strigil- 
lose throughout, or the strigillose pubescence restricted to 
the upper third or half, or (occasionally in Florida) the 
tube completely glabrous or the lower half or two-thirds 
of the tube glabrous and the upper portion variously pubes- 
cent, bearing few to numerous conspicuous pustular glands 
scattered throughout the upper one-third to two-thirds of 
the tube. Calyx-lobes externally usually densely puberulent 
or short-pubescent and hence the margins not appearing 
strikingly ciliate or occasionally (especially in the more 



1975] Amorpha — Wilbur 355 



ms 



merely sparsely puberulent and 



m 



glandular, internally the lobes and occasionally the very 
uppermost portion of the tube densely puberulent to short- 
pubescent (the tube otherwise glabrous within). Adaxial 
calyx-lobes usually triangular-dentate and often narrowly 
so, mostly acute to acuminate but occasionally (most com- 
monly in the Floridian variety with a glabrous calyx-tube) 
oblong and then obtuse, about (0.4)0.5-1(1.2) mm long; 
lateral calyx-lobes broadly to narrowly triangular, mostly 
longer than the adaxial lobes but shorter than the abaxial, 
acute to acuminate and mostly (0.6)0.8-1.2(1.5) mm long; 
the abaxial lobe usually narrowly triangular, acute to 
acuminate and mostly (0.8)1-1.5(2) mm long. Vexillum 
about (4)5-6(7) mm long and (2)2.5-3.5 mm wide, broadly 
obcordate and tapering into a narrow claw, strongly arch- 
ing and incurved laterally, and thus enveloping the inner 
floral parts with an entire to finely erose margin, blue- 
violet, purplish-violet to white. Filaments about 6-8 mm 
long, glabrous. Anthers about (0.3)0.4-0.6(0.8) mm long, 
yellow to yellowish orange. Pistil about 6 mm long with a 
sparsely ciliate or puberulent to glabrous ovary about 1 
mm high, and broadest at or about the middle and tapering 
to either end and with a densely antrorsely pubescent style 
about 5 mm long terminated by a small, truncate, stigmatic 
tip. Fruit about 4-6 mm long and about 1.8-2.5 
broadest at or near the middle and tapering to t 
stipe-like base, conspicuously pustular-punctate in the up- 
per half or two-thirds, densely short-pubescent to glabrous 



mm 



mm 



mm 



laterally compressed, obliquely obovate to very slightly, 
crescent-shaped, the adaxial side straight to slightly arched 
concavely away from the rachis, the abaxial side strongly 
outwardly bowed, terminated by the often 0.3-0.5 mm per- 
sistent base of the style. 

Distribution: Dry, open woods, sandhills, or savannahs, 
typically of the Coastal Plain but occasionally in the Pied- 
mont or mountains from North Carolina south into Florida. 



356 Rhodora [Vol. 77 



This species, the most widespread of the southeastern 
dwarf taxa is not morphologically uniform throughout its 
extensive area. Its variability is most pronounced in 



Florida. 



extreme 



is that element which was formerly designated A. floridana. 
My earlier survey of the southeastern dwarf species 



(Rhodora 56: 261-265. 1954) 



fl 



endemic 



from 



Man 



The suggested differ- 



ent 



by Small (Man. SE. Flora 688. 1933.) to distinguish the 
two taxa is not tenable. The dorsal suture of most fruiting 
specimens of the floridana type is straight, or very nearly 
so, just as in A. herbacea. Only rarely is it as curved as 
shown by Palmer (Jour. Arn. Arb. 12: 194. f. 5. 1931). 
Examination of many more specimens of A. herbacea s.s. 
and of A. floridana has convinced me that the difference 
between these two taxa is merely one of degree of pubes- 
cence. However, there are also a small number of speci- 
mens which seem at least partially intermediate between 
these extremes. Macroscopically they appear much closer 
to extreme A. floridana but microscopically the calyx-tubes 
are pubescent to a varying degree. Some of these tubes 
are sparsely puberulent or short-villous throughout, while 
others are sparingly to densely strigillose throughout. 
These intermediates might indicate introgression from A. 
herbacea, since they are often found in areas in which that 
taxon is known and are often collected with it. Except for 
the apparently isolated area of the glabrous extreme about 
Apalachicola in Franklin County, the range of the so-called 
A. floridana coincides with part of that of A. herbacea in 
western peninsular Florida. I tentatively have distin- 
guished these two less than ideally separable taxa by the 
following key. 



1975] 



Amorpha — Wilbur 357 



KEY TO THE VARIETIES OF A. HERBACEA 



Upper portion of the plant including the leaves usually 
conspicuously pubescent; calyx-tube densely puberulent 
to short-pilose; fruit densely to sparsely puberulent or 
rarely glabrate 3a. A. herbacea var. herbacea. 

Upper portion of the plant and the leaves appearing gla- 
brous or nearly so, or at least not densely pubescent; 
calyx-tube glabrous, sparingly puberulent or sparsely to 

densely minutely strigillose ; fruit glabrous 

3b. A. herbacea var. florid-ana. 



3a A. herbacea var. herbacea, typification : Doubtfully 

present in Walter's Herbarium (bm) as Dr. Carroll 
E. Wood, Jr. did not see a photograph of a specimen 
of this distinctive species in the GH photocopy of 
Walter's Herbarium. A neotype should therefore per- 
haps be designated. 

A rmhe.scens Willd., Berlin Baumz. 17. 1796. 



Michx 



1803. 



(nom. illegit., Intern. Code Art. 63). (typification: 
holotype, P, not seen; phototype, Gh!) 

A. cyanostachya M. A. Curtis, Boston Jour. Nat. Hist. 
1: 140. 1835. typification: North Carolina: Wil- 
mington, M. A. Curtis s.n. (lectotype, ny!). 

A. herbacea var. a typica Schneider, Bot. Gaz. 43: 299. 

1907. 
A. herbacea var. /3 Boyntoni Schneider, Bot. Gaz. 43: 
300. 1907. typification: Florida: PASCO CO., pine 
barrens, Richland, Curtiss 6664 (lectotype, mo!; 
isolectotypes, cu!, ga!, gh !, isc!, min!, neb!, ny!, 
phil!, uc!, us!) 

Usually at least the upper portion of the stem densely 
short-pubescent or puberulent, but occasionally only 
sparsely pubescent and rarely almost glabrous. Rachis of 
leaf usually densely short-pubescent or puberulent through- 



358 Rhodora [Vol. 77 

out or rarely sparsely short-pilose, puberulent or strigillose. 
Rachis of the inflorescence usually densely short-pubescent 
or puberulent. Calyx-tube very densely puberulent or 
short-pilose throughout, and with the pubescence curling 
and spreading and not at all strigillose. Fruit densely to 
sparsely puberulent or short-pubescent throughout 
part glabrous. 



or in 



Typification: Probably not in Walter's Herbarium (bm). 
Supposedly from the area about Walter's plantation in 
present day Berkeley County, South Carolina, where it is 



common. 



Distribution: Dry, open woods, sandhills, or savannahs, 
typically of the Coastal Plain but occasionally in the Pied- 
mont and mountains from North Carolina south into pen- 
insular Florida. (Map 3.) 

Schneider did not treat his two proposed varieties as 
distinct species since they were "apparently connected 
by some intermediate forms." The extremes in leaflet size 
and number are striking but the transition from one 
morphological type to the other is complete. The more 
northern specimens are often separable from those from 

the southern portion of the range, but exceptions are fre- 
quent. 



3b. 



Wilbur 



Mitchell Sci. Soc. 80: 55. 1964. 

A.floridana Rydb., N. Am. Fl. 24: 31. 1919. 



TYPI- 



FICATION: Florida: [no other locality given], 

Chapman s.n. (ny!). 

Upper portion of the stem glabrous or nearly so to 
sparsely short-pilose or puberulent, but occasionally notice- 
ably puberulent or strigillose. Rachis glabrous or nearly 
so to more or less densely puberulent or short-pubescent 
especially above. Rachis of the inflorescence sparsely to 



1975] 



Amorpha — Wilbur 



359 




MAPS 3-5. Map 






Amorpha herbacea var. herbacea. Map 4. 



A. herbacea var. floridana. Map f>. .4. gvorgictna var. gvorgiana (dots 

in the Carolinas and Georgia), A. georgiana var. confusa (triangles) 

and .4. creiiulata (dots with vertical and horizontal lines in southern 

Florida). 



densely puberulent. Calyx-tube glabrous or nearly so to 
sparsely short-puberulent or minutely strigillose through- 
out, and the pubescence, when present, often restricted to 
the upper portion of the tube or at least sparsely dis- 
tributed below, or, if more or less equally dispersed and 
dense, then the pubescence strigillose. 



360 Rhodora [Vol. 77 



Distribution : 



om 



Manat 



lands and fields, sandy roadsides. (Specimens of this spe- 
cies collected by Leavenworth [GH, PH], and others [ny] 
are labeled "East Florida.") (Map 4.) 



4. Amorpha crenulata Rydb., N. Am. Fl. 24: 30. 1919. 

typification : Florida: dade CO., in hummocks, be- 
tween Coconut Grove and Cutler, Small & Wilson 
1898 (holotype, ny!; a presumed isotype, f!). 

A low, apparently rhizomatose shrub about 0.4-1.0(1.5) 
m high. Current season's growth usually dull reddish- 
purple or more rarely olivaceous, usually slender, 1-2(3) 
mm in diameter and but sparsely puberulent to glabrous, 
or very nearly so, with numerous fine longitudinal grooves 
and ridges and sprinkled throughout with few to numerous 
small glandular blistered areas; older branches gray to 
light brownish and less conspicuously lined. Leaves numer- 
ous, spreading to ascendant, (0.8)1.5-2.5(3) dm long. 
Petioles very sparingly puberulent to glabrous and usually 
longer than the breadth of the lowermost leaflet, but oc- 
casionally equaling or exceeded by the breadth of the lower- 
most leaflet, sparingly glandular throughout and densely 
so near the base, (0.3)0.8-1.5(1.8) cm long. Stipules in- 
conspicuous and pigmented like the stem and rachis, gla- 
brous, usually caducous, appressed, setaceous to acicular, 
1.2-2.2 mm long. Rachis slender, about 0.5-1 
eter, glabrous to sparsely puberulent, usually sparingly 
glandular, channeled above. Leaflets (19)23-33(41) in 
number, (0.7)1.2-2.5(4.2) cm long, (2.5) 5-9 ( 
mostly (1.2)2-4(5) times as long as broad, remote, usually 
well separated from the nearest leaflet on the same side of 
the rachis, the interval between those petiolules about (5) 
7-17(20) mm long. Blades of leaflets mostly narrowly to 
broadly oblong or elliptic but varying from almost orbicular 
to ovate-oblong or obovate, with the base commonly broadly 
rounded to almost truncate or gradually tapering and 



mm 



mm 



1975] Amorpha — Wilbur 361 



sometimes acute; the apex broadly rounded to almost trun- 
cate and commonly emarginate or occasionally somewhat 
acute, with the midvein terminating in a sessile or sub- 



mm 



mucro 



Texture of leaflets coriaceous to sub- 
coriaceous when dry, finely reticulate above from the 



mi 



ma 



revolute, usually very noticeably crenate or crenulate. 



M 



veins 



but slightly raised. Both surfaces of the leaflets glabrous 
or rarely very sparsely pubescent beneath along the prin- 
cipal veins; punctate glands on the lower surface usually 
conspicuous and readily apparent to the unaided eye, ap- 
pearing roughly of two sizes, varying from dense and very 
numerous to relatively sparsely scattered and few, ranging 

;ly between 40-180 punctate glands per half-leaflet. 
Petiolule glabrous or rarely sparsely pubescent, pustulate- 
glandular, (1)1.5-2(3) mm long. Stipels acicular, 0.4-1.5 
mm long, glabrous, often not long-persisting and the lower 
often not developing. Racemes solitary or very commonly 
few to several closely associated, slender and elongate, 

ly (0.5)1-2.5(3) dm long and about 0.7-1.8 cm in 



mos 



mos 



diameter, sessile or nearly so, or on a peduncle about 1-2 
(4) cm long, with a conspicuously ridged and grooved gla- 
brous to very sparingly puberulent rachis bearing numer- 



moi 



very sparsely puberulent pedicels (0.5)1-1.8(2.2) mm long, 
subtended by a caducous, gland-dotted, brownish to amber- 
colored, sparsely puberulent, linear-setaceous or narrowly 
spatulate bract nearly 1.2-2.5 mm long. Calyx-tube turbi- 



mo 



mm 



high with the abaxial side slightly longer than the adaxial, 
glabrous or very nearly so throughout, with the few to 
numerous punctate glands mostly restricted to the upper 
half of the tube. Calyx-lobes punctate-glandular, glabrous 
or very nearly so externally with a densely short-ciliate 
margin ; internally the lobes and uppermost portion of the 



362 



Rhodora [Vol. 77 



tube densely matted-puberulent, the lower portion of the 
tube glabrous internally. Adaxial lobes triangular-dentate 
and acute to acuminate to oblong and obtusely rounded and 
about 0.5-1 mm long; lateral lobes broadly to narrowly 
triangular and acute to acuminate and mostly about 0.8- 
1.2 mm long; abaxial lobe usually narrowly triangular and 
acute or more commonly acuminate, mostly (1)1.2-1.8(2) 
mm long. Vexillum about (4.5)5-6(7) mm long and 3-4.5 
mm wide, very broadly obcordate and tapering abruptly to 
a slender claw, strongly arched and incurved and envelop- 
ing the inner floral parts, the margin irregularly finely 
erose to almost entire, white to violet-blue. Filaments about 
6-9 mm long, glabrous. Anthers about 0.5-0.7 mm long, 
yellowish. Pistil about 6 mm long with a glabrous ovary 
about 1 mm high, an antrorsely pubescent style about 5 mm 
long and a .small, capitate, terminal stigma. Fruit about 
4-6.5 mm long and 1.8-2.5 mm wide with the base tapered 
below into a stipe about 1-2 mm long, glabrous, conspicu- 
ously punctate-glandular in the upper two-thirds, exceed- 
ing the calyx-tube by about 2.5-4 mm, laterally compressed, 
obliquely obovate, the adaxial side straight or nearly so, 
the abaxial side strongly outwardly bowed, terminated by 
the often 0.5 mm long, persistent base of the style. 

Distribution : Apparently restricted to Dade County, 
Florida. (Map 5, in part.) 

This appears to be a weakly differentiated species en- 
demic to the southern tip of Florida. It seems very closely 
related to A. herbacea and particularly to the more glabrate 
variant of that species, var. floridana. 



5. Amorpha georgiana Wilbur, Rhodora 56: 261. 1954. 

[10 Jan. 1955] 

A low shrub 0.3-1 m high. Current season's growth 



mm in diameter 



sparsely puberulent or more comm 



numerous 



sprinkled with small punctate glands; the older growth 



1975] Amorpha — Wilbur 36b 



glabrous, dull reddish purple or brownish and less notice- 
ably ridged and occasionally, especially in the southern por- 



moi 



numer 



and then darker and often grayish or black. Leaves 
ous, spreading to ascendant, mostly (3)6-15(18) cm long. 
Petioles glabrous or sparsely to moderately puberulent or 
short-pubescent, ranging in length from practically absent 



more 



mm 



most leaflet, varying from 1-15(20) mm long, sparingly 
glandular to densely .so especially at or near the base. 
Stipules inconspicuous, glabrous, caducous to semipersist- 
ent, appressed, linear-subulate or more commonly setaceous, 
v/ long. Rachis slender, about 

0.5-1 mm in diameter, glabrous to sparingly strigillose, 
puberulent or short-pubescent below and the channeled 
groove similar or rather densely short-puberulent, spar- 
ingly glandular throughout. Leaflets varying from about 
(11)15-43(47) in number, (0.3)0.6-2.5(3.2) 



cm 



(0.2)0.6-1.2(1.6) cm wide, usually about (1)1.5-2.5(3) 
times as long as wide, opposite or nearly so to conspicu- 
ously alternate, approximate and often imbricately over- 
lapping with an interval between petiolules of (0.2)0.3- 
0.5(0.6) cm to rather widely separated with the interval 
between petiolules on the same side of the rachis about 
(0.5)0.8-1.4(1.8) cm long. Blades of the leaflets mostly 
broadly to narrowly oblong or occasionally oblong-elliptic 
to elliptic and rarely ovate-oblong, or the terminal some- 
times obovate, obcordate or even nearly orbicular, with 
the base broadly rounded or almost truncate to subcordate 
and the apex usually obtuse to broadly rounded and com- 



m 



slender mucro about (0.2)0.4-1 mm long and tapering 
gradually to the not at all or rarely very indistinctly swol- 
len, globose or knob-like tip. Texture of leaflets coriaceous 
to subcoriaceous when dried; inconspicuously reticulate 
above from the scarcely elevated venation except for the 
sometimes but slightly depressed midvein; the margin 
usually slightly to conspicuously revolute although occa- 



364 Rhodora [Vol. 77 



sionally flat or nearly so; entire or inconspicuously crenu- 
late and with the midvein prominently and the secondary 
veins slightly elevated beneath; at maturity both surfaces 
glabrous or nearly so, or with the lower surfaces sparsely 
to moderately strigillose, especially along the principal 
veins, or rarely moderately spreading, short-pubescent, the 
punctate glands on the lower surface usually inconspicuous 
but still noticeable to the unaided eye and of uniform size, 
or at least not conspicuously of two size-classes, mostly 
about (30)50-120(160) glands per half-leaflet. Petiolule 
glabrous or -sparsely to densely strigillose, puberulent or 
pilosulose with few to numerous conspicuous to very in- 
conspicuous pustulate glands, 0.7-1.8(2) mm long. Stipels 
acicular or setaceous, glabrous, usually persistent, mostly 
0.8-1.8 mm long. Racemes usually with densely clustered 
flowers or less commonly with the flowers more loosely ar- 
ranged, usually appearing terminally clustered and arising 
from the naked upper portion of the stem or from the axils 
of the upper leaves or occasionally solitary, varying in 
number from (1)4-11(15), mostly (0.2)0.5-2.0(3.0) dm 
long and about (1.0)1.2-1.6 cm in diameter, sessile or 
nearly so or on a peduncle up to about 8 cm long, the rachis 
conspicuously ridged and grooved, glabrous or nearly so 
to sparsely strigillose or puberulent or less commonly 
densely puberulent, sparingly glandular, bearing numerous, 

glabrous to sparingly strigillose or puberulent pedicels 
about 0.4-1 mm long subtended by a caducous, sparingly if 
at all glandular-punctate, sparsely short-pubescent and 
often merely ciliate, brownish to amber-colored, linear to 
linear-subulate bract mostly 1.5-2.5(3) mm long. Calyx- 
tube turbinate, narrowly campanulate or cylindrical, about 
1.7-2.2 mm high with the abaxial side slightly longer than 
the adaxial, glabrous or rarely inconspicuously and very 
sparingly pilosulose, bearing few to numerous more or less 
inconspicuous pustular glands on the upper third to two- 
thirds of the tube. Calyx-lobes externally glabrous to 
puberulent (if the latter, usually sparingly so), and usually 



1975] Amorpha — Wilbur 365 



punctate glandular and with densely white-ciliate margins; 
internally the lobes and occasionally even the uppermost 
portion of the tube densely puberulent to short-pubescent 
(with the tube otherwise glabrous within). Adaxial calyx- 
lobes either triangular-dentate and acute to acuminate or 
oblong, and then obtuse to rounded, mostly 0.4-0.8(1.3) mm 
long; lateral lobes usually longer than the adaxial lobes 
and shorter than the abaxial, usually triangular-dentate 
and acute to acuminate but occasionally oblong and obtuse 
to rounded, mostly (0.6)0.8-1.2(1.5) mm long; abaxial lobe 
usually narrowly triangular to lance-subulate, acute or 
more typically acuminate, mostly (1.0)1.2-1.6(1.8) mm 
long. Vexillum about (4)5-6 mm long and 3-3.5(4) mm 
wide, broadly obovate to obcordate, tapering abruptly into 
a narrow claw, strongly arching and incurving laterally 
and thus enveloping the inner floral parts, strikingly to 
inconspicuously emarginate, the margins entire to very 
finely erose, purplish-blue, bluish or violet in color. Fila- 
ments about 5-8 mm long, glabrous. Anthers about 0.4-0.6 
(0.7) mm long, exserted, yellowish prior to pollen dis- 
charge. Pistil about 6 mm long with a glabrous ovary about 
0.8-1 mm high, including the somewhat stipitate base, 
broadest above the middle, with an antrorsely pubescent 
style about 4-5 mm long terminated by a small, truncate, 
stigmatic tip. Fruit about 4-5.5 mm long and 2-2.5 mm 
wide, broadest at or near the apex and tapering to the 
1-2 mm stipe-like base, glabrous, conspicuously punctate- 
glandular in the upper half or two-thirds, exceeding the 
calyx-tube by about 2.2-3.5 mm, laterally compressed, 
obliquely obovate, the adaxial side straight or nearly so, the 
abaxial side strongly outwardly bowed, usually terminated 
by the 0.2-0.5 mm long, persistent base of the style. 

Distribution: Sandy wire-grass savannahs, pine woods, 
and thickets in the Coastal Plain from southeastern North 
Carolina southward into eastern Georgia. 

This species seems to be represented by two varieties 
which are distinguished in the following key: 



366 Rhodora [Vol. 77 



KEY TO THE VARIETIES OF A. GEORGIANA 

Lateral leaflets (3)6-10(15) mm long and (2)3-5(8) mm 
wide; petioles 1-3(5) mm long; racemes typically solitary 
but rarely with 1-3 short, additional racemes, mostly 

(2)3-5(6) cm long; vexillum reddish-purple 

5a. var. georgiana. 

Lateral leaflets typically (10)15-25(35) mm long and (7) 
9-15(18) mm wide; petioles usually (6)8-15(20) mm 
long; racemes in clusters of (1)3-5(8), 10-20(30) cm 
long; vexillum light to more typically an intense bright 
blue 5b. var. confusa. 

5a. A. georgiana var. georgiana. typification : Georgia : 

TELFAIR CO., sandy wire-grass savannah about 3 
miles northwest of Lumber City, Wilbur 3158 (holo- 
type, gh ! ; isotypes, duke !, ga !, mich !, mo !, nsc !, 

ny!, us!). 

Distribution: Sandy flats bordering streams and low- 
land woods from the central Coastal Plain of North Caro- 
lina south into central coastal Georgia. (Map 5, in part.) 

5b. A. georgiana var. confusa Wilbur, Jour. Elisha Mitchell 

Sci. Soc. 80: 58. 1964. typification: North Caro- 
lina: Brunswick CO., grassy savannah, 7 miles south- 
west of Wilmington, Godfrey & Shunk 4122 (holo- 
type, gh!; isotype, us!). 

A. glabra in the sense of Boynton, Bot. Gaz. 25: 279. 
1898 and in Small's Fl. SE. U.S. 626. 1903, but 
not of Poiret, Encycl. Meth. Suppl. 1: 330. 1810. 

A. caroliniana in the sense of T. & G. Fl. N. Am. 1 : 
305. 1838 in part; Schneider, Bot. Gaz. 43: 302. 
1907 and 111. Handb. Laubh. 2: 74. 1907; Rydb., 
N. Am. Fl. 24: 29. 1919, but not of Croom, Am. 
Jour. Sci. 25: 74. 1834. 

A. cyanostachya in the sense of Palmer, Jour. Arn. 
Arb. 12: 169. 1931, and of Small, Man. SE. Fl. 
639. 1933, but not of M. A. Curtis, Boston Jour. 
Nat. Hist. 1 : 140. 1835. 



1975] Amorpha — Wilbur 367 



Distribution: Pine woods, sandy ridges, savannahs, and 
sandy roadbanks of southeastern North Carolina (Bladen, 
Brunswick, and Columbus Counties). (Map 5, in part.) 

The confused basis of the previous names applied to this 
taxon has been discussed rather fully in an earlier paper 
(Jour. Elisha Mitchell Sci. Soc. 80: 58-61. 1964) and there 
is no need to repeat that discussion here. Although, I 
remain skeptical that the present arrangement of the 
georglana and confusa taxa will prove to be satisfactory 
when more is learned about them, 1 refrain from further 
nomenclatural shifts at the present time. 1 suspect future 
investigation may well demonstrate that the two taxa are 

rwprlv imnressed bv the 



am 



specifically distinct. Perhaps 1 
strikingly different colors of their petals. 



6. Amorpha canescens Pursh, Fl. Am. Sept. 467. 1814. 

Typification : Not seen. Perhaps in Pursh's col- 
lection but Nuttall (Gen. N. Am. PI. 2: 92. 1818) 
states that "Mr. Pursh places his usual mark of v. v. 
to this species, although he has never seen a flower- 
ing specimen except in my herbarium." "On the 



Missouri and M 



Q 



[Wisconsin River] to the Mississippi; around St. 



M 



probably to the Mountains," according to Nuttall. 
A. canescens Nutt., Fraser's Cat. no. 4. 1813. nom. 



nud. 

A. canescens var. (3 leptostachya A. Gray, Mem. Am. 

Acad. II. 4: 31. 1849. nom. nud. 
A. canescens var. glabrata A. Gray, Smithson. Contr. 

Knowl. 3 (Art. 5, PI. Wright): 49. 1852. typifi- 



cation : 



W 



A. canescens var. « typica C. K. Schneid, Bot. Gaz. 

43: 300. 1907. 
A.brachycarpa Palmer, Jour. Arnold Arb. 12: 171. 
1981. TYPIFICATION: Missouri: STONE CO., dry 



368 



Rhodora [Vol. 77 



slopes and ledges, rocky hills near Galena, E. J. 
Palmer 19197 (holotype, a!; isotypes, kanu!, 
min!, us!). 

A. canescens f. glabrata (A. Gray) Fassett, Rhodora 
38: 191. 1936. 



A low, ascendant to erect, apparently rhizomatose, usu- 
ally canescent, shrub mostly about (3)5-8(10) dm high. 
Current season's growth light brown to olivaceous and 
slender (about (1)2-3 mm in diameter) and typically very 
densely grayish tomentose or occasionally but moderately 
so or very rarely glabrous or nearly so, with numerous very 
fine longitudinal striations and apparently eglandular or at 
most with very few minute, amber-colored, very incon- 
spicuous glands ; older branches mostly light gray to brown- 
ish and very slightly, if at all, lined, glabrous or very 
nearly so. Leaves numerous, spreading, about (3.0)6.0- 
12.0(15.0) cm long. Petioles whitish to pale gray, moder- 
ately to very densely tangled tomentose, villous or even- 
spreading short-pubescent or very rarely moderately pu- 
berulent to spreading short-pilose, sometimes equaling but 
usually very much shorter than the breadth of the lower- 
most leaflet, apparently always lacking pustulate glands, 
mostly (0.5)1-3(5) mm long. Stipules inconspicuous, ca- 
ducous, densely canescent-pilose externally, glabrous, dark 
reddish and amber-colored within, initially appressed but 
spreading and often becoming reflexed with age, linear- 
lanceolate, linear to setaceous, mostly (1.2)2-3(3.5) mm 
long, pustular glands apparently always lacking. Rachis of 
leaf slender, mostly about (0.2)0.5-1.0(1.2) mm in diam- 
eter, usually very densely grayish, tangled-tomentose but 
occasionally varying to sparingly so to even strigillose or 
spreading puberulent, channeled above, pustular glands 
apparently always absent. Leaflets mostly (11)27-41(47) 
in number, usually about (0.3)1.0-1.8(2.5) cm long and 
(2.0)4.0-7.0(12.0) mm wide, usually about (1.8)2.0-3.0 
(3.8) times as long as broad, typically crowded and com- 
monly overlapping one another, the interval between peti- 



1975] Amorpha — Wilbur 369 

olules on the same side of the rachis characteristically 
about (2.0)3.0-8.0(10.0) mm long. Blades of leaflets mostly 
ovate-oblong, oblong-elliptic, oblong, elliptic, or even ovate 
with the terminal rarely obcordate and those of the dwarf 
suckers often almost suborbicular to obovate, with the base 
obtuse to broadly rounded, and more rarely subcordate or 

somewhat truncate and the apex obtuse to broadly 



even 



ema 



mu 



m a l 



to (rarely slightly 
swollen at tip on a few leaflets) usually about (0.2)0.4- 
0.8(1.2) mm long. Texture of leaflet subcoriaceous to 
thick-membranous when dry, venation often completely 
obscured or nearly so by the dense pubescence, but when 
discernible usually only finely and inconspicuously reticu- 
late above from the slightly elevated secondary and tertiary 
venation (the midvein somewhat depressed) ; the 
flat to very slightly revolute and entire; the midvein 
markedly elevated beneath and the secondary and tertiary 
venation often forming an inconspicuous net (which is 
usually completely obscured or nearly so by the dense pu- 
bescence) ; both surfaces usually very densely grayish 
tomentose and then appearing canescent to less commonly 
moderately short-pubescent or very rarely nearly glabrous 
with pubescence restricted to the midvein and sometimes 
principal veins and to the margin, and the leaves then 
greenish (the pubescence usually both denser and longer 
below than above) ; pustular-punctate glands usually pres- 
ent beneath, although completely obscured from view by 
the tangled pubescence, when not hidden by pubescence still 
relatively inconspicuous, light olive-green or brown in color, 

m size, very small. Petiolules very 
densely spreading pilose to short-pubescent but rarely 
sparingly short-pubescent, usually appearing canescent, 

(0.5)0.8-1.2 mm long, usually lacking punctate 

glands or glands, if present, very inconspicuous. Stipels 
acicular, about (0.5)0.8-1.2(1.5) mm long, mostly slightly 
exceeding but sometimes almost twice as long as the peti- 
olule, glabrous to sparsely short-pubescent, long-persistent, 



more oi 



s 
mos 



370 Rhodora [Vol. 77 



dark reddish brown, the lower rarely developing beyond 
the swollen base. Racemes usually numerous in the axils 
of the uppermost leaves and often forming a dense com- 
pound cluster, mostly (1)5-20(30) or even more in number. 



r flowered, mostly (2.0)7.0-11 
cm thick; peduncle short, m 



cm 



cm 



— 7 v w 

spreading-pilose to short-pubescent, sparingly and very 
inconspicuously glandular-punctate and bearing numerous, 
very densely arranged, moderately to (more typically) 
densely spreading, wavy, short-pubescent or pilose pedicels 
appearing short and stout with a truncate apex, about 
(0.4)0.6-1.2(1.5) mm long, subtended by a caducous, 
sparsely glandular-pustulate, amber-colored, moderately to 
densely pilose externally and glabrous within, linear- 
setaceous to narrowly lanceolate (with a long tapering 
apex) bract mostly (2.5)3.0-4.0(4.5) mm long. Calyx-tube 
turbinate to obconic-campanulate, mostly (1.5)1.8-2(2.5) 
mm high, slightly asymmetrical with the abaxial slightly 
longer than the adaxial, usually very densely canescent 
throughout, but very rarely sparingly pilose to almost 
glabrous (the tricl 

pilose), usually rather inconspicuously (due to the pubes- 
cence) punctate-glandular, internally glabrous except for 
possibly the uppermost fringe. Calyx-lobes all narrowly 
triangular-lanceolate, obscurely punctate-glandular and 
acute; adaxial lobes (0.6)1-1.4(1.6) 
(1.0)1.2-1.5(1.8) mm long; abaxial lobe (1.2)1.5-2.2(2.5) 



omes m 



mm 



mm 



short- to long-pilose 



more 



um 



about 4.5-6.0 mm long and 2.5-4 mm wide, very broadly 
obcordate and abruntlv tanerinff into a slpnnW rla w fahrmf 



mm 



som 



nate but occasionally also obscurely apiculate, its margin 
entire to irregularly erose, bright violet. Filaments golden- 
brownish to purplish, about 6-8 mm long, glabrous, united 
into a tube for the lowest 1.5-2.0 mm. Anthers about 0.4- 



1975] 



Amorpha — Wilbur 



371 




MAP 6. Amorpha canescens 



mm 



Pistil about 5.0- 



mm lonir with densely pilose ovary about 1 mn 



mm 



densely antrorsely pubescent style about 4-6 

a small, capitate, terminal stigma. Fruit about (3.0)3.5- 

4 5(5.0) mm long and (1.5)1.8-2.2 mm wide, tapering to 



mo 



I, J I \_, Cj vy I l X v> T T A * *- V *-" +s w ■*■ V-* ^** ■» — — — — — — W v 

villous-canescent but very rarely glabrous, punctate-glandu- 
lar in the upper half or two-thirds, exceeding the calyx- 

_ — fill 1 _ — 



mm 



so and the abaxial side strongly outwardly bowed, termi- 



mm 



372 Rhodora [Vol. 



i i 



Distribution: Roadsides, fields, prairies, hillsides, and 
open woodlands from Indiana west into the Dakotas, Wyo- 
ming, and New Mexico, and southern Canada south into 
eastern Texas. (Map 6.) 

The reasons for recognizing neither A. canescens var. 
glabrata A. Gray nor A. brachycarpa have been presented 
in an earlier paper (Jour. Elisha Mitchell Sci. Soc. 80: 
62-63. 1964) and will not be repeated here. 

An apparently rarely encountered hybrid between A. 
canescens and A. fruticosa may be better discussed here 
than elsewhere since the overall aspect of the plant is such 
that it most likely would be taken as an aberrant, oversized 
A. canescens. The rarity of the supposed hybrid certainly 
attests to the effectiveness of the isolating mechanisms 
between the two species for they occupy the same general 
geographical area over a very extensive region of the 
central United States. 

Palmer (Rhodora 55: 158-159. 1953) described the 
hybrid of these two species as Amorplta X notha from a 
single plant found on a "low rocky bank between upland 
prairie and alluvial valley of Center Creek, about 1 mile 
north of Webb City, Jasper Co., Missouri." Both species 
are common in southwestern Missouri and yet Palmer, who 
was extremely familiar with the plants of that area and 
who had a particular interest in Amorpha, knew only one 
example of the hybrid. In addition to the two collection 
numbers made by Palmer from the type plant (Palmer 
52047, 27 May 1951, in flower; isotypes, F, mo, SMU, UMO) ; 
and Palmer 52930, 9 August 1951, in fruit (F, umo). 
Palmer made an additional collection (#56605) from the 
same area, if not from the same plant, two years later. 
Another collection which I take to represent this hybrid 
was collected by Palmer, #25305, 4 June 1924: dry rocky 
ledges, limestone bluffs of Spring River, near Melugin, 
Jasper Co., Missouri (a, min, umo) and was cited, mis- 
takenly, I believe, (Jour. Arnold Arb. 12: 167. 1931) as 
A. canescens var. glabrata A. Gray. 



1975] Amorpha — Wilbur 37 



o 



The hybrid differs from .4. canescens with which it has 
the greatest overall affinity in its taller habit, much .shorter 
calyx-lobes and particularly in its dorsally strongly curved 
fruit. From its A. fruticosa parentage, it differs in densely 
canescent, pilosulose calyx-tube as well as the finely canes- 
cent leaves and young stems. The practice of giving bi- 
nomials to hybrids of this sort has, I believe, very little to 
recommend itself as a botanical custom. 



mici 



7. Amorpha nana Nutt., Fraser's Cat. 1813. TYPIFICATION : 

"Collected near the Mandan towns, 1600 miles up 
the Missourie," Nuttall, not seen. 

ophylla Pursh. Fl. Am. Sept. 466. 1814. 
now. illegit., Art, 63. typification : not seen; 
Pursh stated that he based name on a specimen 
presented to him by Meriwether Lewis of the 
Lewis and Clark Expedition. 
A. punctata Raf., New Fl. 3: 14. 1837. typifica- 
tion: Specimen not seen. "Discovered by Brad- 
bury in the upper Missouri." 

A low, erect, apparently rhizomatose shrub aobut 3-6 
(9) dm high. Branches of the current season light reddish- 
brown to pale olivaceous; clustered near the top of the 
stem; slender (about 1-2(3) mm in diameter), sparsely 
or, more typically, moderately strigillose but bec< 
glabrate below with age and with numerous very fine, 
longitudinal grooves and ridges and a sparse to moderate 
sprinkling of small inconspicuous punctate-pustular, amber- 
colored glands; older branches usually light gray or pale 
brownish, only slightly, if at all, lined. Leaves numerous, 
spreading to ascendant, (1.5)3-7(10) cm long. Petioles 



mi 



somew 



puberulent, becoming glabrous or nearly so, equaling or 
usually longer than the breadth of the lowermost leaflet 
(rarely the breadth of leaflet is greater than length of 
petiole), about (2)4-8(10) mm long, very sparingly pustu- 
late-glandular. Stipules inconspicuous, pigmented like the 



374 Rhodora [Vol. 77 



young stem and petiole, caducous, short-pubescent along 
margins and outer surface and often with a moderately 
long tuft of trichomes near tip, appressed to spreading, 
setaceous to linear, (2)3-5(6.5) mm long, occasionally 
sparingly pustulate-glandular. Rachis of leaf very slender, 
mostly about 0.3-0.7 mm in diameter, glabrous or nearly 
so to densely very short-puberulent or strigillose and 



m 



channeled above. Leaflets mostly (7)13-27(31) in number, 
(2)6-13(18) mm long, (2)3-6(7.5) mm wide, usually (1) 
1.8-2.9(3.5) times as long as broad, rarely overlapping to 
more commonly remote, usually the interval between peti- 
olules on the same .side of the rachis about (3)4-7(9) mm 



long. 



m 



som 



or obovate to almost orbicular, with the base usually 
rounded or occasionally with margins gradually tapering 
and with the base then cuneate, the apex usually broadly 
rounded or even truncate and commonly emarginate but 
occasionally acute, with the midvein exserted into a slender, 
tapering mucro usually about (0.6)0.8-1.2(1.5) 
Texture of leaflet coriaceous to subcoriaceous when dry, 
finely and inconspicuously reticulate above from the some- 
what elevated secondary and tertiary venation (the mid- 



mm 



vein 



margin flat or more 



somen 



ulate ; the midvein conspicuously elevated beneath and with 
the secondary and sometimes the tertiary veins somewhat 
elevated and sometimes forming an inconspicuous net; 



mi 



the margin usually sparingly short-pubescent, puberulent 
or somewhat strigillose or rarely glabrous or nearly so; 
the punctate glands conspicuous and readily visible to the 
unaided eye on the lower surface, appearing to be of uni- 
form size or at least not falling mostly into two more or 
less distinct size classes, usually with about (10)20-35(55) 
punctate glands per half-leaflet. Petiolule densely to spar- 
ingly short-pubescent or strigillose and sometimes becoming 



1975] Amorpha — Wilbur 375 

glabrate with usually few to several small pustulate glands 
and mostly about (0.5)0.7-1 mm long. Stipels acicular, 
about (1.0)1.5-2.5(3) mm long, often 2 times or more as 
long as the petiolule, glabrous, long-persistent. Racemes 
solitary at the tips of the current season's growth, con- 
tracted and very densely flowered, mostly (2.0)3-7(9) cm 
long and about 1-1.5 cm in diameter, subsessile or the 
peduncle about 0.3-1.5(2.0) cm long; rachis conspicuously 
ridged and grooved, moderately to very densely puberulent 



m 



densely clustered, moderately to densely puberulent or 
strigillose (often becoming sparsely so or even glabrate in 
fruit), often long-persistent (and with 



some 



what expanded apex), about 1.0-2.0(2.5) mm long, sub- 
tended by a caducous, sparsely glandular-pustulate, brown- 
ish to amber-colored, moderately to densely puberulent, 
linear-setaceous to narrowly spatulate bract mostly (2.5) 
.°>.0-4.0(5.0) mm long with a long, tapering apex. Calyx- 
tube turbinate, mostly 1.8-2.2 mm high with the abaxial 



side slightly longer than the adaxial, glabrous throughout 



more 



arranged in rows and restricted to the upper half of the 
tube. Calyx-lobes all narrowly triangular-lanceolate and 
acute, punctate-glandular, glabrous or very sparsely pu- 
berulent externally but with densely short-ciliate margins; 



l 



most 



tube densely matted short-puberulent and the lower portion 
of the tube internally (as well as externally) glabrous; 
adaxial lobes about (0.8)1.0-1.5(1.8) mm long; lateral lobes 
mostly (1.0)1.2-1.8(2.0) mm long; abaxial lobe (1.5)1.8- 
2.0(2.2) mm long. Vexillum about 4.5-6.0 mm long and 
.•5.5-4.5 mm wide, very broadly obcordate, abruptly taper- 
ing into a slender claw, strongly arched and incurved and 
enveloping the inner floral parts, its margin emarginate 
and usually finely erose, dark purple. Filaments purplish, 
about 6-8 mm long, glabrous; anthers about 0.4-0.6 mm 
long, purplish. Pistil about 4.0-6.0 mm long with a gla- 
brous ovary about 0.8 mm high, an antrorsely pubescent 



376 



Rhodora 



[Vol. 77 




MAP 7. Amorpha nana. 



1975] Amorpha — Wilbur " 377 



mm loner and a small, capitate, terminal 



stigma. Fruit about 4.5-5.5 mm long and 2.0-2.8 mm 



mm 



long, glabrous, conspicuously punctate-glandular in the 
upper two-thirds, exceeding the calyx-tube by about 2.0- 



mm 



nearly so, the abaxial 
side strongly outwardly bowed, terminated by persistent 



mm 



Distribution : 



from 



Iowa and Minnesota west into the Dakotas and Colorado 
and from southern Manitoba and Saskatchewan south into 
New Mexico; primarily a plant of the northern plains. 
(Map 7.) 



8. 



lorpha glabra Poir., Encycl, Meth. Suppl. 1: 330. 1810. 
typification : type not seen, perhaps at P; based on 
a specimen grown in cultivation in Paris. 

?A. glabra Desf., Tabl. Ecole Bot. Paris 192. 1804, 
nom. nud. 

?A. glabra Pers., Syn. PI. 2 : 295. 1807. nom semi- 



nud. 



Biltmore 



typification: North Carolina: rocky woods, Bilt- 
more, Biltmore Herb. 14 (lectotype, us!; fruiting 
specimen collected 29 August 1896 on sheet num- 
bered 331562; isolectotypes, a!, cu!, GH !, min!, 
mo!, ND-Greene!, ny!, RM !, us!). 

?A. fruticosa var. glabra Bean, Trees and Shrubs 
Brit. Isles 1 : 193. 1914. 

A much-branched, erect shrub mostly 1-2 m high. Cur- 
rent season's growth light to dark purplish or light-brown 
to olivaceous, slender to moderately thickened, about 1-2 
(3) mm in diameter, usually completely glabrous or at most 
very sparsely and obscurely strigillose to puberulent, spar- 
ingly, if at all, beset with inconspicuous, minute, amber- 
colored pustulate glands; branches of the preceding year 



378 Rhodora [Vol. 77 



mostly grayish to blackish. Leaves wide-spreading - , mostly 
(1.0)1.4-2.2(3.0) dm long. Petioles mostly (1.6)2.0-4.0 
(5.7) cm long, typically much longer than the width of the 
lowermost leaflet, glabrous or very nearly so or very spar- 
ingly puberulent, sparsely beset, if at all, with amber-col- 
ored pustulate-glands. Stipules inconspicuous, appressed 
when present, caducous, thin, sparsely to 



mo 



marg 



dark reddish-brown, typically linear to linear-lanceolate, 
about 2.5-4.5 mm long, apparently eglandular. Rachis of 
leaf about 0.5-1.0 mm in diameter, usually greenish-oliva- 
ceous, glabrous or rarely at most exceedingly sparingly 
puberulent or strigillose, usually sparingly glandular pus- 
tulate throughout, interval between petiolules on the same 
side of the rachis about (0.8)1.5-3.0(3.6) cm long. Leaflets 
mostly (7)11-15(21), either opposite or alternate, about 
(1.4)2.2-4.6(7.4) cm long and (1.0)1.4-2.8(3.6) cm wide, 
mostly (1.2)1.4-2.2(2.7) times as long as broad, character- 
istically widely spaced and not overlapping, usually broadly 
oblong, oblong-elliptic to elliptic or less commonly ovate- 
oblong to ovate or rarely nearly orbicular, basally typically 
very broadly rounded to truncate or nearly so or even 
subcordate but rarely gradually tapering to even acute, 
apically very broadly rounded to strongly obtuse and almost 
invariably conspicuously emarginate. Midvein usually ter- 
minating in a swollen, globose, glandular tip either sessile 
or exserted on a stalk about 0.2-0.4 mm long. Both surfaces 
with venation scarcely either elevated or depressed on 
either surface except for the conspicuously elevated mid- 
vein and the primary and secondary veins obscurely reticu- 
late, glabrous or rarely sparsely and inconspicuously stri- 
gillose beneath along the midvein and principal lateral 
veins, usually conspicuously glandular-punctate; punctate 
glands variable in size, usually numerous, mostly ranging 
in number from about (30)50-150(300) glands per half- 
leaflet. Margin entire, inconspicuously and finely undulate 
to conspicuously crenulate, slightly revolute to more or less 
flush-margined. Petiolules about 2.0-4.0(5.5) mm long, 



1975] Amorpha — Wilbur 379 

usually glabrous but occasionally sparingly to moderately 
puberulent or strigillose, mostly sparsely glandular-pustu- 
ate, wrinkled when dried. Stipels acicular to setaceous, 
mostly 1.2-2.0 mm long (but the tips quickly caducous and 
thus appearing shorter), glabrous, usually long-persistent, 
dark reddish-brown. Racemes often solitary and terminal 
or few and terminally clustered, 1-3(4) in number, with 
densely clustered flowers, mostly about (0.5)1.0-1.8(2.8) 



im 



with a peduncle 1-6 cm long; rachis sparsely glandular- 
punctate, glabrous or very sparingly strigillose or puberu- 
lent. Pedicels glabrous or at most sparsely strigillose or 
puberulent, usually eglandular, about (0.8)1.0-2.5(3.0) mm 
long; bracts caducous, apparently eglandular, golden- 
brown, usually puberulent or ciliate, linear-oblong or linear- 
lanceolate, about (0.5)0.8-1.5(2.0) mm long. Calyx-tube 
broadly turbinate or campanulate, mostly (2.0)2.2-3.0 
(3.2) mm high, somewhat asymmetrical with the abaxial 
side slightly higher than the adaxial, glabrous externally 
and internally throughout excepting the densely short- 
ciliate rim, bearing 1 or 2 rows of small, inconspicuous, 
pustulate-glands mostly restricted to the upper third of the 
tube. Calyx-lobes often obsolete or nearly so and the top 
of the calyx then appearing truncate or the lobes low and 
depressed with the calyx-rim appearing weakly undulate 
but when noticeably developed usually broader than high, 
mostly rounded or broadly triangular-dentate; adaxial lobes 
often rounded and shorter than the lateral lobes but fre- 
quently triangular-dentate and then occasionally longer 
than the lateral lobes, when developed mostly (0.1)0.2-0.3 
(0.5) mm long; lateral lobes often broadly rounded but 
commonly broadly triangular-dentate and when developed 
beyond mere undulations then mostly (0.1)0.2-0.4(0.5) mm 
long; adaxial lobe typically triangular and acute although 
occasionally rounded, mostly (0.1)0.2-0.6(0.8) mm long. 
Vexillum about (4.5)6.0-8.0(8.5) mm long and 4.0-6.0 mm 
wide, broadly obcordate, abruptly to gradually tapering 



mm 



380 



Rhodora 



[Vol. 77 




MAPS 8-10. Map 8. Amorpha glabra. Map 9. A. schweriniL 
Map 10. A. paniculata. 



usually moderately to strongly emarginate, bright reddish- 
purple. Filaments yellowish to golden, about 8-11 mm long, 
glabrous; the monadelphous tube noticeably exserted at 
maturity from the calyx-tube for about (0.5)1.0-2.0 mm, 
the united portion of the filaments mostly about 3.0-4.5 mm 
long; anthers about 0.5-1.0 mm long, golden-yellow to 
orange. Pistil about 7.0-9.0 mm long with a glabrous, obo- 
vate ovary about 1 mm high and an antrorsely short- 
pubescent style terminated by a truncate stigmatic tip. 
Fruit about (6.5)7.5-9.0(10.5) mm long and (2.5)3.0- 



1975] Amorpha — Wilbur 381 



4.0(4.5) mm wide, broadest at or above the middle and 
tapering- to the slender, conical base, glabrous throughout, 
the upper two-thirds sparsely to moderately covered with 
small but usually conspicuously pustulate glands, obliquely 
obovate with the adaxial valve straight or nearly so and 
the abaxial valve very strongly outwardly bowed above, 



mm 



the style. 

Distribution : Endemic to the southern Appalachian Mts. 
of western N. and S. Carolina, northeastern Ga. and east- 
ern Tennessee. (Map 8.) 

The attribution of this species to Arkansas and Okla- 
homa (Jour. Arnold Arb. 12: 174-175. 1931, and Proc. 
Oklahoma Acad. Sci. 27: 69. 1947.) is, I believe, an error. 
Specimens so annotated in the past seem to me to be A. 
ouachitensis whose closest relative appears to be the central 
Texan A. roermeriana (=A. texana) and not the southern 
Appalachian endemic. 

9. Amorpha schwerinii Schneider, Bot. Gaz. 43: 301. 1907. 

[as Schwerini]. typification : North Carolina: 
ROWAN CO., Dunn's Mountain, alt, 1200 ft. /. K. 
Small s.n. (holotype, mo!; isotypes, f! ny!). 

A. densi flora F. E. Boynton ex Small, Fl. SE. U.S. ed. 
2 1342. 1913. TYPIFICATION: North Carolina: 



GASTON CO., slopes of Crowder's Mountain, Bilt- 
more Herb. 14756b (lectotype, ny!; isolectotypes, 

min! ny! rm!). 

A bushy, widely branched, erect shrub mostly (1)1.5- 
2.5 m high. Current season's growth dull olive-brown to 
reddish-brown or dull grayish to almost black and typically 



mm 



moder 



small 



pustular glands; older branches dark grayish to blackish 
but becoming dark reddish-brown with numerous, incon- 
spicuous longitudinal fissures and more or less orbicular, 



382 



Rhodora [Vol. 77 



becoming 



Buds globose to ovoid, scales densely puberulent to short- 
pilose. Leaves wide-spreading, about (0.5)0.8-1.2(2.2) dm 
long. Petioles mostly (0.7)1.0-1.8(2.6) cm long, mostly 
equaling or exceeding the length of the lowermost leaflet 
but occasionally shorter, moderately to densely pubescent 
with spreading hyaline trichomes (up to 0.7 mm long), 
bearing none or but few pustular, amber-colored glands. 
Stipules inconspicuous, usually appressed while present but 
soon caducous, copiously tawny puberulent to curly short- 
pubescent externally and glabrous or nearly so within, dark 
reddish-brown, typically linear to linear-lanceolate, about 
3.0-4.5 mm long, eglandular. Rachis of leaves slender, 
mostly 0.5-0.7 mm in diameter, typically brownish to gray- 
ish, moderately to densely puberulent to spreading short- 
pubescent with the trichomes up to about 0.5 mm long, 
eglandular or the pustulate glands sparse and inconspicu- 



ous. 



mostlv (7)19-27(29) in numbe 



(0.5)1.5-3.0(4.0) cm long and (0.4)0.8-1.5(1.9) cm wide, 
typically about (1.3)2.0-3.0(3.8) times as long as broad, 
rarely overlapping, the interval between petiolules on the 
same side of the rachis mostly about (0.6)0.8-1.5(1.8) cm 
long; usually narrowly to broadly oblong but varying from 
elliptic to elliptic-oblong and rarely from ovate-oblong to 
ovate or even almost orbicular in unusual terminal leaflets, 
typically with the base broadly rounded but varying to 
somewhat truncate or even slightly subcordate and the apex 
mostly obtuse to broadly rounded or occasionally emar- 
ginate; venation slightly elevated above and more pro- 
nouncedly so below. Midvein usually swollen terminally, 
usually exserted for 0.2-0.5(0.8) mm. Margin entire to very 
obscurely crenulate and inconspicuously revolute. Lower 
surface densely pilose with soft ascendant or spreading hy- 
aline, tapering trichomes mostly 0.3-0.6 mm long particu- 
larly pubescent along the midvein, moderately to densely 
beset with olivaceous to amber, punctate glands beneath 
(about 30-120 glands/half-leaflet) ; upper surface minutely 
and obscurely but copiously short-pubescent with hyaline, 



1975] Amorpha — Wilbur 383 



appressed to spreading - trichomes about 0.1-0.3 mm long. 
Petiolules usually (0.5)1-2 mm long, densely spreading 
pilosulose to short-pubescent with hyaline trichomes mostly 
0.2-0.6 mm long, indistinctly glandular-pustulate. Stipels 
acicular, about 1.0-2.2 mm long, often equaling the petio- 
lules in length, typically long-persistent, dark reddish-brown, 
pilosulose and often densely so at base and glabrous or 
nearly so above, sparingly pustulate. Racemes 1 or some- 
times 2 and the principal one usually terminating strong 
shoots and the second when present smaller and arising at 
or near base of the terminal cluster, mostly (2.0)4.0-8.0(12.0) 
cm long and 1.0-1.5(1.8) cm in diameter; rachis of inflores- 
cence densely pilosulose to short-pubescent, eglandular or 
nearly so. Pedicels puberulent to pilosulose, eglandular, 
about (0.8)1.0-1.5(2.0) mm long; bracts caducous, sparingly 
glandular-pustulate, moderately to densely pilosulose ex- 
ternally and glabrous within, linear to linear-lanceolate, 
about 2.0-3.5 mm long. Calyx-tube turbinate to obconic- 
cylindrical, mostly 1.8-2.5 mm long, somewhat asym- 
metrical with the adaxial side slightly longer than the 
abaxial, externally sparingly to densely spreading pilosulose 
throughout, moderately to densely beset with amber-col- 
ored, pustulate glands in the upper one-fifth to one-third 
but eglandular below. Calyx-lobes mostly linear to linear- 
lanceolate (very rarely the four paired lobes triangular- 
dentate), acute, sparingly to moderately glandular-punctate, 
short-pilose both externally and internally, mostly equaling 
or exceeding the length of the tube. Adaxial lobes about 
(1.2)2.0-3.0 mm long; lateral lobes about (1.5)2.0-3.2 mm 
long; abaxial lobe (1.8)2.5-3.5 mm long. Vexillum about 
4.5-6.5 mm long, 3.0-4.2 mm wide, very broadly obovate to 
obcordate, tapering into the slender claw of about 1.5 mm 
long, apically emarginate and sometimes also inconspicu- 
ously apiculate, margin entire to irregularly erose, purplish. 
Filaments yellowish to golden, about 6-8 mm long, glabrous, 
united at base into a tube 1.5-2.0 mm long; anthers about 
0.5-0.8 mm long, golden-yellow. Pistil about 5-7 mm long 
with a densely antrorsely pubescent style about 4-6 mm 



384 



Rhodora [Vol. 77 



stigma. Fruit about 5-6.5 mm 



1.8-2.2 mm 



pubescent or rarely glabrous or nearly so, usually conspicu- 
ously glandular-pustulate in upper half, the adaxial valve 
straight or nearly so and the abaxial very strongly out- 



mm 



the style. 

Distribution : River banks and open slopes from the inner 
North Carolina Piedmont southwest into Alabama. (Map 9.) 

This is morphologically one of the most distinctive and 
clearcut species within the genus. 



10. Amorpha paniculata T. & G., Fl. N. Am. 1: 306. 1838. 

typification : Texas: T. Driimmond 461 (lectotype, 



ny!; isolectotype, gh!). 



m 



relatively unbranched. Current season's growth usually 
densely puberulent to spreading short-pubescent or glabrate 
in age, often somewhat canescent and lacking pustulate 
glands. Buds obovoid, densely appressed short-pubescent, 
canescent. Leaves wide-spreading, mostly (1)2-4 dm long. 
Petioles (2)3-6(9) cm long, usually noticeably longer than 
the width of the lowermost leaflet, stout, canescent, densely 
puberulent. Stipules caducous, setaceous, pubescent, about 
3 mm long. Rachis of leaf about 2.5 mm in diameter, 



more 



m 



elliptic, usually (1.5)3-6(8) cm long and 1.5-3(5.3) cm 
wide, typically (1.4)1.8-2.2(2.6) times as long as wide, 
opposite or more commonly alternate, usually symmetrical, 
typically broadly rounded both apically and basally, con- 
spicuously reticulate below with the veins elevated often to 
the fourth degree of branching. Midvein exserted up to 



mar 



ing but rarely swollen. 



model 



to densely covered with spreading, crisped, hyaline to 
tawny, pilosulose trichomes but rarely sparingly to mod- 



19 75] Amorpha — Wilbur 38 ° 

erately short-pubescent or even glabrate; upper surface 
glabrate or sparingly short-pubescent to puberulent espe- 
cially along the principal veins, usually inconspicuously 
and sparingly to moderately beset beneath with small pus- 
tulate, punctate glands. Petiolules (0.2)0.4-1.0 cm long, 



moi 



somewhat canescent, sparingly and inconspicuously pustu- 
late-glandular. Racemes several to numerous, (l)o-l- in 
number, about (0.5)1.5-3.0(4.0) dm long; rachis of inflor- 
escence finely pilosulose, canescent. Pedicels slender, about 
1-2 mm long, pilosulose; bracts very slender, setaceous, 
pilosulose, about 2-2.5 mm long. Calyx-tube funnel-form 



mm lone, moder 



omes 



to 



e rivaling lhwiwih^o ^ „ ^ 

to moderately and inconspicuously glandular-pustulate 
above Calyx-lobes inconspicuously glandular-pustulate, 
densely but finely pilosulose; adaxial lobes broadly to nar- 
rowly triangular-dentate or ovate, acute, 0.8-1.0(1.-) mm 
long; lateral lobes longer and usually more narrowly lanceo- 
late, acute to acuminate, (1.0)1.2-1.4 mm long; abaxial 
lobe (1 2)1.5-2.0 mm long, linear to linear-lanceolate. Yex- 
illum 5-7 mm long, 3-4 mm wide, entire to slightly erose 
apically, purple. Filaments about 5-6 mm long united for 
about half their length, glabrous ; anthers about 0.6-0.8 mm 
long, orange. Style densely villous; ovary glabrous. Fruit 
4-6(8) mm long and 2-2.5 mm wide, broadest somewhat 
above the middle, tapering to the base, with the adaxia 



mar 



the abaxial margin strongly outwardly bowed glabrous, 
conspicuously glandular-pustulate. Seed reddish-brown, 
about 3.5 mm long and 1.8 mm wide, smooth. 

Distribution : Thickets, bogs, swampy woods and ditches 
of southwestern Arkansas, west central Louisiana and 
eastern Texas. (Map 10.) 



m 



synonyms 



The only syno- 



Cies IS suggesicu vj iw *--- » 

nym sometimes associated with it has been A. roemenana 



386 Rhodora [Vol. 77 



Scheele, but I believe this binomial properly belongs to the 

central Texan plant previously referred to as A. texana 
Buckl. 



11. Amorpha nitens Boynton, Biltmore Bot. Stud. 1: 139. 

1902. typification : Georgia: swamp near Waynes- 
boro, Biltmore Herbarium s.n. (holotype: not seen, 
designated as at the Biltmore Herbarium and hence 
presumably at the us). 

A. nitens var. Jeucodermis E. J. Palmer, Jour. Arnold 
Arb. 12: 177. 1931. typification: Georgia: 
thicket along lake, Augusta, Boynton 7035 (holo- 



type, Gh!). 



An erect, branching shrub 1-3 m tall. Current season's 
growth usually blackening or at least darkening upon dry- 
ing, glabrous to sparingly or even moderately puberulent 
to pilosulose, usually eglandular or bearing only a few 
scattered, small inconspicuous glands. Buds compressed 
and broadly oblong or ovoid, blackish when dried, sparingly 
pilosulose to glabrate except for the shortly puberulent 
margins of the outer scales. Leaves more or less wide- 
spreading to ascendant, mostly 1-1.8(2.2) dm long, usually 
conspicuously darkening or blackening upon drying. Peti- 
oles glabrous to moderately crispy short-pilose, eglandular 
or with a few, inconspicuous, small, pustulate glands, typi- 
cally longer than the width of the lowermost leaflet, mostly 
(1.5)2-3.5(5) cm long. Stipules caducous, linear to linear- 
setaceous, eglandular, drying blackish, usually glabrous 
except externally with an apical tuft of tawny pubescence, 
mostly 3-5 mm long. Rachis of leaf about 0.6-1.0 mm in 
diameter, glabrous to moderately crispy-pilosulose, usually 
eglandular but rarely very sparingly and inconspicuously 
glandular. Leaflets (7)9-15(19) in number, oblong to el- 
liptic-oblong or occasionally somewhat ovate, mostly 2-4(7) 
cm long and (1.0)1.5-3.5(4.5) cm wide, typically 1.5-2.5 



times as long as wide, opposite or more typically alter- 
nate, usually widely spaced, symmetrical, usually broadly 



1975] 



Amorpha — Wilbur 387 



moi 



rounded and often emarginate, entire; secondary venation 
usually little, if at all, elevated beneath. 



M 



shortly exserted and but little tapered. Lower surface 
of leaflets glabrous to moderately spreading pilosulose 

restricted to the 



imes 



mm 



midvein, eglandular or inconspicuously and often rather 
sparingly beset with small punctate glands; upper surface 
eglandular and glabrous and often shiny at maturity but 
occasionally sparsely beset with extremely minute and 
fine, appressed short-pubescence. Petiolules about 2-6 .1 
long glabrous to moderately spreading pilosulose, eglandu- 
lar to inconspicuously and very sparingly pustulate gland- 
ular. Racemes erect, solitary or with several clustered 
together, (0.5)0.8-1.5(2.5) dm long; rachis of inflorescence 
glabrous to moderately pilosulose or puberulent, eglandular 
or very sparingly and inconspicuously pustulate-glandular. 
Pedicels 1-2.2 mm long, glabrous to moderately pilosulose 
or crispy pubescent, slender, eglandular, blackening upon 
drying; bracts usually caducous, basally attached on pedi- 
cel or distally as much as midway towards the tip, very 
narrowly linear to setaceous, 1.5-3(4) mm long, glabrous 
to externally tangled tawny-pilosulose especially along the 
margins, usually eglandular but occasionally inconspicu- 
ously and very sparingly pustulate glandular, typically 
blackening upon drying. Calyx-tube slenderly obconic to 
broadly funnelform to rarely even roundedly campanulate, 
mostly 2-2.5 mm long, glabrous to moderately pilosulose, 
often completely eglandular but occasionally very sparingly 
beset with inconspicuous, minute pustular glands in the 
upper third. Calyx-lobes with a ciliate fringe and with the 
outer surface glabrous to pilosulose, eglandular; adaxial 
calyx-lobes usually broadly rounded and obtuse but occa- 



mm 



more 



triangular, about 0.4-1.0 mm long; abaxial lobe triangular 
and acute and often narrowly so, 0.6-1.4 mm long. Vexil- 
lum 4.5-6 mm long, 3-4.8 mm wide, broadly obovate, grad- 



388 Rhodora [Vol. 77 



mm 



slightly to strongly arched, apically broadly rounded and 
entire to irregularly erose, apparently reddish purple 
["blue-purple fide Palmer"] and with the claw and blade 
both enveloping the filaments. Filaments 5-7 mm long, 
united into a sheath for the lower 2-2.5 mm, glabrous; 



mm long, yellow. Style moderately 
J mm long; ovary glabrous. Fruit 

) mm wide, broadest ahnvp tha mir 



mm 



tapering basally with the adaxial suture straight or slightly 
curved and the abaxial strongly outwardly bowed, glabrous, 
moderately pustulate glandular distally. Seeds smooth, 
dark reddish-brown to blackish, 3-4.2 mm long. 

^ Distribution. Thickets and moist woods along creeks, 
rivers and bottomlands from southern Illinois southeast 
into Georgia and west into Louisiana and eastern Okla- 
homa. (Map 11.) 



12. Amorpha laevigata Nutt. in Torr. & Gray, Fl. N. Am. 

1 : 306. 1838. typification : "Banks of the Arkan- 
sas, near Salt River," Nuttall s.n. (holotype, BM, not 
seen; phototype: a!) 

A. laevigata var. typica Schneider, Illustr. Handb. 
Laubh. 2:74. 1907. 

Shrub 1-2(3) m tall. Current season's growth sparingly to 
moderately strigillose and glandular-pustulate, light brown 
to more characteristically deep reddish-purple. Buds sub- 
orbicular to ovoid, compressed; the scales sparingly short- 
pubescent. Leaves wide-spreading to moderately ascendant, 
mostly (0.5)0.8-1.5(2.4) dm long. Petioles (0.6)1.0-2.5 
(3.4) cm long, about 1 mm in diameter, very sparingly 
strigillose or glabrous, sparingly to moderately glandular- 
pustulate, usually longer than the width of the lowermost 
leaflet. Stipules caducous, linear-subulate to setaceous, 



of leaf 0.5-1 mm in diameter 



mm long. Rachis 



1975] 



Amorpha — Wilbur 



389 




MAPS 11-14. Map 11. Amorpha nitevs. Map 12. A. laevigata. 
Map 13. A. roemeriana. Map. 14. A. ouackitensis. 

strigillose, sparingly to moderately glandular-pustulate. 
Leaflets (9)13-19(25), oblong or oblong-elliptic to rarely 
obovate, usually (1.0)1.6-3.2(4.2) cm long and (0.4)1.0- 
1.5(2.2) cm wide, typically (1.5)1.8-2.4(2.8) times as long 
as wide, usually alternate and symmetrical, characteristi- 
cally basally rounded but rarely acutely tapering, apically 
usually broadly rounded and often emarginate; venation 
little elevated beneath except for the mid vein ; margin 
entire to slightly crenulate. 



M 



mor 



moi 



390 Rhodora [Vol. 77 



mm. Lower surface glabrous or very sparingly strigillose 
especially along the midvein, moderately to densely beset 
with conspicuous punctate glands often appearing to be of 
two size classes or rarely nearly epunctate; upper surface 
glabrous. Petiolules (1)2-3(4) mm long, glabrous or more 
typically sparingly to moderately strigillose, conspicuously 
and densely beset with markedly elevated pustulate glands ; 
stipels usually persisting, dark reddish or blackish, about 
1-2 mm long. Racemes solitary or loosely clustered, 1-3(8), 
about (0.5)1.0-2.0(3.0) dm long; rachis of inflorescence 
very sparingly strigillose, eglandular or sparingly glandu- 
lar-pustulate. Pedicels usually 0.6-1.2 mm long and spar- 
ingly to moderately strigillose, eglandular; bracts caducous, 
setaceous to narrowly lanceolate with a long-tapering apex, 
sparingly to moderately strigillose, 1.5-3 mm long, glandu- 
lar-pustulate. Calyx-tube funnelform to somewhat cam- 
panulate, about 1.5-3 mm high, glabrous or sparingly to 
moderately strigillose, moderately to more typically densely 
beset with pustulate glands in upper two-thirds. Calyx- 
lobes sparingly to densely strigillose with a dense fringe of 
white trichomes, glandular-pustulate; adaxial lobes broadly 
rounded to triangular-dentate and acute, about 0.2-0.5 mm 
long; lateral lobes usually triangular dentate and acute to 
acuminate but occasionally broadly rounded, 0.5-0.8 mm 
long; adaxial lobe narrowly triangular-dentate, 0.8-1.2 mm 
long. Vexillum about 4-6 mm long and about 4.0-4.5 mm 
wide, entire to finely erose, bright blue to deep violet blue. 
Filaments to 9 mm long, united for about half their length, 
glabrous; anthers about 0.5-0.7 mm long, orange. Ovary 
glabrous; style antrorsely pubescent. Fruit about 4.5-6 
long and 2-2.5 mm wide, with the adaxial suture straight 
or slightly incurved near the apex and the abaxial suture 
strongly outwardly bowed, glabrous or rarely sparingly 
strigillose, conspicuously glandular-pustulate on the distal 
three-fourths. 

Distribution : A rare species of prairies, open woods and 
creek banks of eastern Texas and Oklahoma. (Map 12.) 



mm 



1975] 



Amorpha — Wilbur 391 



13. Amorpha roemeriana Scheele, Linnaea 21: 461. 1848. 

typification : "In margine rivulorum prope, Aus- 
tin," F. Roemer s.n. (holotype: not seen by me or by 
anyone else who has commented upon it.) 

A. fruticosa var. [1] subglabra A. Gray, Boston 
Jour. Nat. Hist. 6: 174. 1850. typification: 

Texas: on a creek near Fredericksburg, June 1817, 
Lindheimer s.n. (holotype, oh!). It is doubtful if 
Gray intended or actually did publish this varietal 
name. The type-style is different than that em- 
ployed in this paper for varieties and its punctua- 
tion is also different. The next entry in the 
Plantae Lindheimerianae is also listed by Gray 

as A. fruHcosa var. mb glabra. 
A. laevigata var. pubescens A. Gray, Smithsonian 
Contr. Knowledge 3: 49. 1852. typification: 
''Eastern" Texas, Charles Wright s.n. (holotype, 

gh!). 

A.texana Buckley, Proc. Acad. Nat. Sci. Philadel- 
phia 1861: 452. 1862. TYPIFICATION: Texas: "On 
the Pierdenalis River," S. B. Buckley s.n. (holo- 
type, PHIL.!). 

A.subglabra (A. Gray) Heller, Contr. Herb. Frank- 
lin & Marshall Coll. 1: 48. 1895. 
A. texana [var.] mollis Boynton, Biltmore Bot. Stud. 



1: 139. 1902. A needlessly created "nom. nor. 
based on "A. laevigata pubescens A. Gray." 

A. laevigata var. pubescens f. mollis (Boynton) C. 
K. Schneider, Bot. Gaz. 43: 307. 1907. 

A. texana var. glabrescens E. J. Palmer, Jour. Arn- 
old Arb. 12: 180. 1931. typification: Texas: 
[comal CO.] Comanche Spring; New Braunfels 
etc., Lindheimer 743 (holotype, a!; isotypes, ARiz!, 

f!, gh!, mo!, ny!, okl!, phil!, uc!, us!). 

Shrub 1-3 m tall. Current season's growth sparingly to 
densely puberulent or strigillose or rarely glabrous, occa- 



392 Rhodora [Vol. 77 



pustulate. Buds ovoid, comp 



moderately 



what keeled, sparingly to moderately puberulent. Leaves 
strongly divergent, (0.5)1-1.5(2) dm long. Petioles (0.5) 
1.5-3.0 cm long, about 1 mm in diameter, very sparingly to 
occasionally densely puberulent or strigillose, sparingly to 
moderately glandular-pustulate, usually equaling or longer 
than the width of the lowermost leaflet. Stipules caducous 
and rarely seen, linear, tawny puberulent, ca. 2 mm long. 
Rachis of leaf slender, 1-1.6 mm in diameter or less, very 



mo 



moi 



densely puberulent. Leaflets (7)9-11(15), oblong or 
typically broadly oblong to oblong-elliptic or rarely obovate 
to suborbicular, usually (1.0)2.5-4.0(5.2) cm long and 
(0.7)1.5-2.5(3.8) cm wide, typically (1.0)1.3-2.9(3.0) times 
as long as wide, opposite or more commonly alternate, 
typically symmetrical, usually broadly rounded basally and 
apically either broadly rounded or more characteristically 
emarginate and often conspicuously so; venation little ele- 
vated beneath other than the midvein and to a slight degree 
the secondary veins; margins entire to conspicuously cren- 
ulate. Midvein rarely exserted as much as 0.8 mm long as 
a tapering mucro but more typically terminating in a 
slightly swollen knob at the margin or tapering less than 
mm beyond. Lower surface usually sparingly to mod- 
erately pilosulose or puberulent with fine, hyaline trichomes 
or rarely glabrous or nearly so, usually inconspicuously 
and moderately beset with small punctate glands; upper 
surface glabrous or at least glabrate to moderately pilosu- 
lose or finely puberulent especially along the principal 
veins. Petiolules usually 2-5(7) mm long, glabrous to 
densely puberulent or pilosulose, often becoming wrinkled 
upon drying, usually sparingly to moderately glandular- 
pustulate but rarely eglandular. Stipels inconspicuous, 
often caducous, dark reddish, about 1.2-2 mm long. Ra- 
cemes solitary or loosely clustered, 1-3(6) in number, about 
(4)6-12(20) cm long; rachis usually sparingly to moder- 
ately strigillose or puberulent, rarely densely tawny or 



1975] Amorpha — Wilbur ?>93 



hyaline puberulent or short-pubescent, sparingly glandular- 
is 1-1.5 mm long, usually sparingly 



mos 



to moderately puberulent or strigillose; bracts caducous, 
setaceous to narrowly lanceolate and then often with a 
long-tapering apex, puberulent or pilosulose, 1.2-2.2 mm 
long. Calyx-tube funnelform, usually 2.5-3,8 mm long, very 
sparingly to moderately strigillose to pilosulose with hya- 
line trichomes, moderately glandular-pustulate in upper 
third; lobes moderately to more characteristically densely 
short-pubescent above with a conspicuous fringe of white 
trichomes, glandular-pustulate; adaxial lobes broadly 
rounded to broadly triangular-dentate and acute, (0.2) 
0.4-0.6(0.8) mm long; lateral lobes broadly rounded to more 
characteristically acutely triangular-dentate, 0.5-0.8(1.0) 
mm long; abaxial lobe usually narrowly triangular-dentate, 
about (0.8)1.0-1.2(1.4) mm long. Vexillum about 5-7 mm 
long, 5-6 mm wide, emarginate and occasionally apiculate, 
entire to slightly erose, purple. Filaments at maturity to 
10 mm long, united for up to two-thirds their length, gla- 
brous ; anthers about 0.6-0.8 mm long. Style villous ; ovary 
glabrous. Fruit about 6-7 mm long and 2.5-3.5 mm wide, 
plump at maturity, with the adaxial margin straight or 



somew 



m 



margin strongly outwardly bowed, glabrous to 
strigillose, conspicuously glandular-pustulate above. Seed 
bright brown, about 3 mm long and 2 mm wide. 

Distribution : All creek bed and stream banks of the "Hill 
Country" of central Texas in the eastern portion of the 
Edwards Plateau. (Map 13.) 

There is no record in the literature that anyone has ex- 
amined authentic material of A. roemermna Scheele. A. 
Gray stated (Boston Jour. Nat. Hist. 6: 175. 1850.) that 
A. roemermna was "doubtless a form of A. fruticosa or of 
A. particulate." Later Gray (Proc. Acad. Nat. Sci. Phil. 
1862: 162) indicated that A. roemeriana was apparently 
the same as A. texana Buckl. which he synonymized with 
A. laevigata var. pubescens. Watson (Smithsonian Misc. 



394 



Rhodora [Vol. 77 



Coll. 258. 188. 1878), Schneider (Bot. Gaz. 43: 307. 1907) 
Rydberg (N. Am. Fl. 24: 27. 1919) and Palmer (Jour. 
Arnold Arb. 12: 179. 1931) have all listed Scheele's A. 
roemeriana as a synonym of A. paniculata T. & G. 

Unfortunately for the sake of stability, it would seem 
most unlikely that A. roemeriana can be considered a 
synonym of A. paniculata. A. paniculata is a largely East 
Texas species of thickets and low swampy or marshy 
ground. A. roemeriana was originally collected along a 
creek bend near Austin which is its eastern range limit. 
Amorpha texana has been collected in Travis Co. while 
A. paniculata is not known from the vicinity. The original 
description itself is far more suggestive of the central 
Texan species than it is of more easternly ranging A. 
paniculata. I have no doubt that A. roemeriana is an 
earlier name for the species recently known as A. texana. 

Palmer (1931) recognized a glabrous or subglabrate and 
a spreading pubescent variety of this species. There seems 
to be no geographic segregation of these pubescence types 
and actually there seems to be as much of a continuum in 
vestiture as one could expect in so small a number of col- 
lections as are available of this species. I concur with 
Turner (The Legumes of Texas, p. 143, 1959) and Correll 
and Johnston (Man. Vase. PI. Tex., p. 818, 1970) who did 
not recognize the varieties either. Both of these taxonomic 
works suggest that A. laevigata is not specifically distinct 
from A. texana, Turner suggesting that A. laevigata "per- 
haps is best treated as a narrow-leaved eastern variety of 
A. texana" while Correll and Johnston indicated that it was 
"perhaps only a form of A. texana." I believe the differ- 
ences in petal color, calyx shape, texture and glandularity 

etc. all indicate considerable morphological divergence be- 
tween the two taxa. 



14. Amorpha ouachitensis Wilbur, sp. nov. TYPIFICATION : 

Oklahoma: LE FLORE CO., dry, rocky (sandstone) 
slopes of Black Fork Mountain, near Page, Palmer 
20572 (holotype, a!; isotypes, ga!, ny!, us!). 



1975] Amorpha — Wilbur 395 

Frutex erectus 1-2 m altus. Foliola (7)9-13(17), 
lateralia oblonga vel oblongo-elliptica, opposita, basi 
et apice rotundata vel emarginata, (1.2)2.5-4.0(7.0) 
cmlongaet (0.9)1.5-2.5(3.6) cm lata. Tubus calycis 
2.8-3.2 mm altus, ± glaber, glanduloso-punctatus 
in quarta parte superiore. Lobi calycis glabri vel 
pubescentes; lobi abaxiales 0.4-0.6 mm longi; lobi 
laterales 0.5-0.7 mm longi; lobi adaxiales 0.6-0.9 



(1.2) mm 



mm 



eum 



Shrub 1-2 m tall. Current season's growth glabrous or 



more 



sparingly pustulate glandular with inconspicuous lenticular 
glands. Buds ovoid to almost globose with the scales mod- 
erately appressed pubescent on the outer surface and api- 
cally densely ciliate with tawny-villous trichomas. Leaves 
strongly divergent, 7-23 cm long. Petioles 1.2-2(2.4) cm 
long and about 0.5-1.5(2) mm in diameter, m 



mo 



ichomes, 
ly equal- 
ling or longer than the width of the lowermost leaflet. 
Stipules caducous, lanceolate, densely tawny villous api- 
cally, ca. 3-4 mm long including the apical tuft but only 

Rachis of 

leaf mostly 1 mm in diameter or less, very sparingly 
pilosulose to even glabrous, sparingly glandular-pustulate. 
Leaflets (7)9-13(17), typically broadly oblong to oblong- 
elliptic or oblong, mostly (1.2)2.5-4.0(7.0) cm long and 



mm 



(0.9)1.5-2.5(3.6) cm wide, typically (1.4)1.5-2(2.4) times 
as long as wide, usually opposite, typically symmetrical, 
usually broadly rounded basally and apically character- 
istically conspicuously emarginate to broadly rounded; 
venation but little elevated beneath other than the mid- 
vein; margins entire to inconspicuously undulate-crenate. 
Midvein exserted either as a tapering mucro about 0.2- 
0.6(1.0) mm or only slightly exserted and terminating in a 
swollen knob. Lower surface of the leaflets glabrous or 



396 



Rhodora [Vol. 77 



m 



comm 



cent with fine, hyaline to pale tawny, slender trichomes 
and usually conspicuously beset with numerous pustulate, 
amber-colored glands; upper surface glabrous or glabrate 
to moderately pilosulose with inconspicuous, slender tri- 
chomes and lacking pustulate glands. Petiolules 
(4) mm long, glabrous or glabrate to moderately short- 
pubescent or pilosulose, sparingly to moderately glandular- 
pustulate or rarely eglandular. Stipels inconspicuous, some- 
times caducous, drying dull dark reddish-brown, mostly 



mos 



mm 



Racemes solitary or in loose clusters of 2-4, mostly (8) 



cm 



or even moderately spreading short-pubescent with hyaline 
or very rarely tawny trichomes and eglandular or very 
sparingly and inconspicuously pustulate-glandular; pedi- 
cels about 1-1.5 mm long, glabrous to sparingly puberulent; 
bracts caducous, narrowly lance-oblong, about 1-1.5 mm 
long, marginally short-pubescent with stiff, hyaline tri- 
chomes. Calyx-tube narrowly conical to funnelform, about 
_.8-3.2 mm long, usually glabrous or nearly so to moder- 
ately puberulent or spreading short-pubescent, sparingly 
glandular pustulate in the upper quarter; lobes usually 
glabrous but occasionally moderately appressed pubescent 
on the external surface and densely margined with a con- 
spicuous fringe of white trichomes, eglandular to sparingly 
pustulate glandular; adaxial lobes broadly rounded, mostly 
0.4-0.6 mm long; lateral lobes broadly rounded or perhaps 
more characteristically broadly triangular-dentate, about 
0.5-0.7 mm long; abaxial lobe narrowly triangular-dentate 
with an acute to acuminate apex, about 0.6-0.9(1.2) mm 
long. Vexillum about 5-7.5 mm long and 4.5-6 mm wide, 
strongly emarginate but otherwise entire, purple. Fila- 
ments at maturity 8-10 mm long, united for up to 3/5 their 
length, glabrous; anthers about 0.6-0.8 mm long. Style 
antrorsely pilosulose; ovary usually glabrous. Fruit about 
7-9 mm loner and 3-4 mm wide and with the adaxial margin 



1975] Amorpha — Wilbur 397 

either straight or more typically mostly straight but with 
the upper portion bent sharply abaxially and the abaxial 
margin strongly outwardly bowed, glabrous or occasionally 
somewhat appressed, short-pubescent and conspicuously 
pustulate-glandular. Seed solitary, dark brownish to black- 
ish, about 4-5 mm long and 2.5-3 mm wide. 

Distribution: Ouachita Mountains of west-central Arkan- 
sas and southeastern Oklahoma. (Map 14.) 

This species has previously been included within the 
concept of other species. Palmer cited specimens of it as 
A. glabra Poir. (Jour. Arnold Arb. 12: 175. 1931) and also 
A. virgata (Jour. Arnold Arb. 12: 182. 1931) which I have 
treated as a synonym of A. fruticosa. Amorpha glabra is 
in my opinion a southern Appalachian endemic while the 
range of A. fruticosa s.lat. spans the continent. I believe 
the morphology of A. ouachitensis indicates a closer rela- 
tionship with the rare central Texan endemic .4. roemeriana 
(= A. texana) than with either A. glabra or A. fruticosa. 

Representative Specimens: Arkansas: conway CO., rocky moun- 
tainsides, Petit Jean Mt., Vemaree MTKH (SMU) ; GARLAND CO., shah' 
outcrop on banks of Ouachita River near Hot Springs, Palmer 24252 

(a); logan co., rocky glades, top of Magazine Mt., Palmer 24187 

(A, MO, UAKK); MONTGOMERY CO., Mount Ida, open hanks of Ouachita 

River below bridge, Demaree 37965 (smu) ; polk CO., Rich Mt., rich 
woods E of lodge near Lover's Leap, Tucker 12285 (Dtkk). Okla- 
homa: lk FLORE CO., Ouachita National Forest, Goodman 2545 (on, 
isc, MO, NY. OKL), rocky slopes of Blackfort Mt., near Page, Palmer 
2001.3 (a); MCCURTAIN Co.: rocky sandy soil in woods 37 mi. N of 
Broken Bow, Stratum. 1146 (OKL) ; PUSHMATAHA CO., rocky stream 
bank, 4 mi W of Albion, Stratum 41C.1 (ll, okl). 



15. Amorpha fruticosa L., Sp. PL 2: 713. 1758. 

A. fruticosa a vulgaris Pursh, Fl. Am. Sept. 2: 466. 

1814. 
A. fruticosa p emarginata Pursh, Fl. Am. Sept. 2: 

466. 1814. 
A. fruticosa y angustifolia Pursh, Fl. Am. Sept. 2: 

466. 1814. 



398 



Rhodora [Vol. 77 



A. croceolanata Wats., Dendr. Brit. 2: t. 139. 1825 

[as crocealanata] • 
A. emarginata (Pursh) Sweet, Hort. Brit. 121. 1827. 
A. frag vans Sweet, Brit. PL Gard. 3: t. 2U. 1828. 
A. caroliniana Croom, Amer. Jour. Sci. 25: 74. 1834. 

typification : North Carolina: near New Bern, 

Loomis s.n. (lectotype, NY!). 
A. fmticosa var. 5 coerulea Loudon, Arb. Brit. 607. 

1838. 
A.humilis Tausch, Flora 21: 750. 1838. typifica- 
tion : undetermined ; based on cultivated plants 

grown in Prague. 
A. fruticosa var. 4 Leivisii Loudon, Arb. Brit. 2: 607. 

1838. 
A. tennessensis Shuttle worth ex Kunze in Delect. 

Sem. Hort. Lips. 1848: 1. 1848; Linnaea 24: 191. 

1851. typification: Tennessee: Ad rivulos prope 

Dandridge, Rug el s.n., June 1842 (isolectotype, 

GH!). 
A. fmticosa var. caroliniana (Croom) S. Wats., 

Smithsonian Misc. Coll. 258. 188. 1878. 
A. fruticosa var. crispa Kirchn., Arb. Muse. 370. 

1864. 
A.pendula Carr., Rev. Hort. 1870-71: 378. 
A. fruticosa var. pendula (Carr.) Dipp., Laubh. 3: 



691. 189 



o. 



A. fruticosa f. albiflora Sheldon, Bull. Geol. & Nat. 
Hist. Surv. Minnesota 9: 72. 1894. typification: 
Minnesota: MILLE LACS CO., on the banks of the 
Rum River, 3 miles north of Milaca, Sheldon 
s.n. (holotype, min!). 

A.virgata Small, Bull. Torrey Club 21: 17. pi. 171. 

1894. typification: Georgia: de kalb CO., NW. 
slope of Stone Mountain, alt. 1000-1400 ft., Snmll 
s.n., 3 July 1893 (lectotype, ny!; isolectotypes, 

ga!, gh!, isc!, min!, mo!, uc!, us!). 

A. fruticosa var. croceolanata (Wats.) Mouillefort, 
Arb. Arbriss. 1 : 577. 1894. 



1975] Amorpha — Wilbur 399 



A. angustifolia (Pursh) Boynton, Biltmore Bot. 

Stud. 1: 139. 1902. 
A.fruticosa f. aureo-variegata Schwerin, Mitteil. 

Deutsch Dendr. Ges. 16: 255. 1907. 
A.fruticosa var. typica Schneider, Bot. Gaz. 43: 

304. 1907. 
A. fruticosa f. crisptt (Kirchn.) C. K. Schneider, 

Illustr. Hand. Laubh. 2: 72. 1907. 
A.fruticosa f. pendula (Carr.) C. K. Schneider, 

Illustr. Handb. Laubh. 2: 73. 1907. 
A.fruticosa var. humilis (Tausch) Schneider, Bot. 

Gaz. 43:305. 1907. 

A. occidentalis Abrams, N. Y. Bot. Gard. Bull. 6: 

394. 1910. typification : California: San Diego 

River, near Old San Diego Mission, Abrams 3425 

(holotype, DS, not seen; isotypes, a!, ds!, GH !, mo!, 

ny!, pom!, vc\, us!) 

A.Bushii Rydb., N. Am. PI. 24: 31. 1919. typifica- 
tion: Florida: uncommon near spring, Chatta- 
hoochee River, Bush 13 (holotype, ny!; isotype: 

a!) 
A.Curtissii Rydb., N. Am. PL 24: 30. 1919. typifi- 
cation: Florida: duval CO., low ground near 
river, Jacksonville, Curtiss 4703 (holotype, ny!; 

isotypes, kanu!, us!) 

A.arizonica Rydb., N. Am. Fl. 24: 33. 1919. typi- 
fication: Arizona: COCHISE CO., along streams, 
Ramsey Canyon, Huachuca Mountains, Gooding 
136 (holotype, ny!; isotypes: ariz!, GH !, neb!, 
ny!, rm!, uc!). 

A. occidentalis var. arizonica (Rydb.) E. J. Palmer, 
Jour. Arnold Arb. 12: 185. 1931. 

A.fruticosa f. humilis (Tausch) E. J. Palmer, Jour. 
Arnold Arb. 12:189. 1931. 

A.fruticosa f. coerulea (Loud.) E. J. Palmer, Jour. 
Arnold Arb. 12: 189. 1931. 



400 



Rhodora [Vol. 77 



A. fruticosa var. angustifolia f. glabrata E. J. 
Palmer, Jour. Arnold Arb. 12: 191. 1931. TYPIFI- 
cation: Texas: brazos CO., shores of lake, Kurten, 
Palmer 13479 (holotype: a!; isotype, mo!). 

A. fruticosa var. tennesseensis (Shuttleworth ex 
Kunze) E. J. Palmer, Jour. Arnold Arb. 12: 192. 
1931. 

A. fruticosa var. oblongifolia E. J. Palmer, Jour. 
Arnold Arb. 12: 192. 1931. typification : Arkan- 
sas: Phillips CO., low alluvial ground, along Mis- 
sissippi River, near Helena, Palmer 26628 (holo- 
type, a!; isotypes, ds!, mo!, uark!). 

A. occidentalis var. emarginata E. J. Palmer, Jour. 
Arnold Arb. 12: 185. 1931. TYPIFICATION: Ari- 
zona: [maricopa CO.] Fish Creek, Apache Trail, 
Eastwood 8745 (holotype, a!; isotype, CAs!). 

A. emarginata Eastwood, Proc. Calif. Acad. Sci. 20: 
148. 1931, not A. emarginata (Pursh) Sweet, 

1827. typification: Arizona: [maricopa CO.] 

Fish Creek, Apache Trail, Eastwood 8745 (holo- 
type, cas!; isotype: a!). 

A. DeWinkeleri Small, Man. Se. Fl. 689. 1933. 
typification: Florida: lee CO., prairies near Ft. 
Shackleford, Big Cypress, Small 8349 (lectotype, 
ny!; isolectotypes, gh!, mo!, us!). 

A. fruticosa var. angustifolia f. latior Fassett, Rho- 
dora 38: 190. 1936. typification: Wisconsin: 
ST. croix CO., stony shore of Lake St. Croix, 10 
miles south of Hudson, Fassett 17014 (holotype, 
wis, not seen; isotypes, gh !, mo!, ny!). 

A. fruticosa var. occidentalis (Abrams) Kearney & 

Peebles, Jour. Washington Acad. Sci. 29: 483. 
1939. 

An erect shrub (1)2-3(4) m tall with one to several 
stems arising from the base and often branching above 
sufficiently so as to appear bush-topped. Current season's 
growth rarely glabrous or densely to moderately pilosulose 



1975] Amorpha — Wilbur 401 

to puberulent or strigillose, occasionally becoming glabrate; 
eglandular or very sparingly and inconspicuously pustulate- 
glandular, grayish to dark reddish-brown, occasionally 
somewhat glaucescent. Buds ovoid to subglobose, light 
brownish to dark reddish-brown, usually more or less gla- 
brous except for the ciliate margins of the scales or occa- 
sionally pilosulose. Leaves wide-spreading to somewhat 



mos 



• -ciiuemi- .uvouv - - long ' llsuallv remaining green 

bTtTcc^Tionally" turning brownish upon drying but not 
blackening Petioles moderately puberulent to crispy pilo- 
sulose and often becoming glabrate. rarely glabrous, eg- 
landular or sparingly pustulate with small, inconspicuous 
glands, typically longer than the width of lowermost leaflet, 
mostly 1-1 cm long. Stipules caducous, narrowly linear, 
eglandular or very sparingly and inconspicuously pustulate, 
reddish-brown, externally and especially apically tawny to 
hyaline pilosulose, usually 2-4 mm long. Rachis of leaf 
about 0.5-1 mm in diameter, rarely glabrous or more typi- 
cally sparingly to densely puberulent to crispy pilosulose 
or glabrate, eglandular or very sparingly and inconspicu- 
ously pustulate-glandular. Leaflets 9-21(31). oblong to 
elliptic-oblong or elliptic or even occasionally 0V *te,Jl)£. 
4(5) cm long and (0.5)1-2(2.7) cm 



mos 



„mes as long as wide, usually opposite or subopposite, 
basally symmetrical or rarely asymmetrical, acute to 



emai 



UiOclUlV numucu L/docn.j, *~ ~- — - 

nate apically, entire or nearly so; secondary venation mod- 
erately elevated beneath. Midvein usually slender, exserted, 
about 0.5-1.5 mm long and tapering, but occasionally only 
very shortly exserted and little, if at all, tapering. Lower 
surface of ieaflets sparsely to densely spreading puberulent 
to crispy pilosulose (occasionally so densely so as to appear 
velvety) when young or rarely either glabrous or some- 
times glabrate in fruit, often strigose or strigillose or oc- 
casionally even short-spiculate with the trichomes either 
hyaline or ashy or even tawny; eglandular or inconspicu- 
ously beset with small pustulate glands or very rarely 
rather conspicuously glandular-punctate; upper surface 



402 Rhodora [Vol. 77 



usually very short-puberulent to glabrous when young and 
typically glabrate when mature but occasionally densely 
pilosulose, eglandular. Petiolules about (1.5)2-4 



mm 



more 



ing pilosulose, occasionally becoming glabrate, or very 
rarely glabrous, usually inconspicuously pustulate-glandu- 



lar. Stipels slenderly cylindric, setaceous mostly 2-4 mm 
long, glabrous to moderately puberulent. Racemes erect, 
solitary or more typically with several to many clustered 
together, usually densely flowered, (0.5)1-2(2.5) dm long; 
rachis of inflorescence usually moderately to densely pu- 
berulent or pilosulose, sometimes becoming glabrate in age, 
eglandular or very sparingly and inconspicuously pustulate- 
glandular. Pedicels mostly 1-2.2 mm long, sparsely to 
densely spreading puberulent or more typically crispy pilo- 
sulose or short-spiculate or even glabrous, eglandular or 
very sparingly and inconspicuously pustulate-glandular; 
bracts caducous, borne basally on the pedicels, very nar- 
rowly linear to setaceous, mostly 1.5-3 mm long, sparingly 
to densely pilosulose externally, usually eglandular, typi- 
cally dull reddish-brown. Calyx-tube usually obconic but 
varying to funnelform or even somewhat campanulate, 
mostly 2-3(4) mm long, glabrous or spiculate, or sparingly 
to densely puberulent or pilosulose, sparsely to moderately 
pustulate-glandular on the upper third of tube with smail 
and inconspicuous to large and conspicuous amber-colored 
glands but occasionally eglandular; lobes with a hyaline, 
ciliate-fringed margin and with the external surface spar- 
ingly to moderately pilosulose and either eglandular or very 
sparingly beset with inconspicuous pustulate glands ; adax- 
ial lobes broadly rounded to broadly triangular-dentate, 
about 0.2-0.5 mm long; lateral lobes usually acute and tri- 
angular-dentate but occasionally broadly rounded, about 
0.3-0.8(1.0) mm long; abaxial lobe narrowly triangular- 
dentate, acute, (0.5)0.8-1.2 mm long. 



um 



mm wi 



tapering to a rather indistinct claw of about 1-1.5 mm long, 
moderately to strongly arched, apically broadly rounded 



1!>7.~>] 



Amorpha — Wilbur 403 

and entire or indistinctly emarginate to irregularly erose 
appearing dark reddish-purple either- fresh or dried and 
with the claw and blade enfolding and enveloping the fila- 
ments and style. Filaments 6-8 mm long, united into a 
sheath for the lower 1-2(3) mm, glabrous; anthers 0.4-0.6 

long, yellow. Style moderately ascending pilose, 5-7 
mm long'; ovary glabrous or rarely pubescent. Fruit 5-9 



mm 



mm long and (2.0)3-4.5 mm 



to strongly curved along the abaxial suture and the abaxial 
suture strongly outwardly bowed, glabrous to densely ap- 
pressed short-pubescent, conspicuously pustulate-glandular 
or eglanclular. Seeds smooth, reddish-brown, 3.5-4.5 mm 



long. 



Distribution: Southern Quebec south into Florida and 
west into northern Mexico, southern California, and Wyo- 



ming. (Map 15.) 



m 



mer 



lengthy synonymy, extremely variable. Ni 
and or varieties have been segregated from it in the past 
and many have been recognized in most recent fioristic 
treatments. The striking variability is such that it might 
seem reassuring indeed to have a formally designated sys- 
tem by which one could categorize the very dissimilar 
appearing plants. Unfortunately, however, the examination 
of many hundreds of specimens soon demonstrates the 
inadequacy of even the numerous described segregates to 
convey the extent of the variability encountered not to 
mention the not inconsiderable number of specimens which 
clearly match the characteristics of one described taxon 
in its older growth and another in its younger and more 
recently formed portions. Clearly then what might be con- 
veniently referred to as the fruticosa-zoi 
tremely diverse assemblage of populations whose variability 
is due both to environmentally induced plasticity and also 
to the presence of a great number of biotypes. I have found 
it taxonomically unclarifying to attempt to categorize this 
variability formally by the recognition of the previously 



ex- 



404 



Rhodora 



[Vol. 77 




013 

C 






>» 



Oh 



1975] Amorpha — Wilbur 



405 



segregated species, varieties and forms and have concluded 
that the variability could best be treated as representing 
one extremely polymorphic taxon. The alternate approach 
is exemplified by Steyermark's treatment in the flora of 
Missouri where as many as four of the five varieties recog- 
nized as occurring in Missouri are known from a single 
county. No indication of ecological preference is suggested 
to account for at least partial isolation of these populations 
and they are in fact stated to be "intergrading." 

TAXA EXCLUDED FROM THE GENUS 
AND BINOMIALS AND TRINOMIALS 

OF UNCERTAIN POSITION 



normals or trinom 



momi 



completeness been included in the following list which per- 
haps could have been excluded since they were not validly 
published. Names published without a diagnosis or descrip- 
tion or without a reference to one that was effectively pub- 
lished are nomina nuda and hence technically could be 
ignored (ICBN Art. 32 (3)). Also binomials or t: 
which were originally published in synonymy are not 
validly published (ICBN Art. 34(4)) and hence could be 
ignored (Art. 12). 

Amorpha arborea Hort. ex Schkuhr, Bot. Handb. Deutschl. 
Gew. 2: 333. 1796. [Publication not seen by me; this 
binomial was reportedly published as a synonym.] 

Amorpha canescens [f.] glabrcsccns Zabel in Beissner et al., 
Handb. Laubh. Ben. 268. 1903. [Publication not seen by 
me; reportedly this bionomial was published as a now. 



m 



N. C. Fassett.] 



Amorpha eolorata Raf., Aut. Bot. 81. 1840. [Appalachian 
Mts.] Rafmesque himself questioningly suggested that it 
might be A. caroliniana Croom, and Merrill (Index 
Rafinesquianus, p. 142. 1949) suggested, again with a 
question mark, its identity with A. nitens Boynton. I 
am unable to identify it with any certainty. 



406 Rhodora [Vol. 77 



Amorpha crocea Hort. ex Lavallee, Arb. Segrez. 60. 1877. 
[Publication not seen by me but this binomial reportedly 
appeared as a nom. nud.] 

Amorpha dealbata Hort. ex Lavallee, Arb. Segrez. 60. 1877. 



me 



] 



Amorpha discolor Raf., Aut. Bot. 79. 1840. [Alabama.] 
Merrill (Index Rafinesquianus, p. 142. 1949.) synony- 
mized this binomial with A. fruticosa L. 

Amorpha elata Hayne, Dendr. Fl. 134. 1822. [Publication 
not seen by me. Rydberg cited this binomial in the syn- 
onymy of A. fruticosa L.] 

Amorpha elatior Hort. ex Lavallee. Arb Rwrw 60 



1877. 



me but this binomial 



edly a nom. nud.] 



Amorpha flexuosa Raf., Aut. Bot. 80. 1840. [Unaka Mts. of 
Carolina.] Perhaps a synonym of A. fruticosa L. as sug- 
gested by Merrill (Index Rafinesquianus, p. 142. 1949). 

Amorpha fruticosa var. frag ana Bean, Trees and Shrubs 



me 



Brit. Isl. 1: 193. 1914. [Publication not seen by 
Palmer states that the brief description would apparently 
exclude it from synonymy of A. fragrans Sweet but that 
its identity is undeterminable.] 

Amorpha gaertnerii Hort. ex K. Koch, Dendr. 1: 70. 1869. 

[Publication not seen by me; reportedly a nom. nud.] 
Amorpha gardnerii Hort. ex. Kirchner, Arb. Muse. 370. 



1864. 
mid.] 



me: renortedlv a nom 



Amorpha glandulosa Blanco, Flora de Filipinas. ed. 1. 555. 
1837. [Philippines.] = Dakiea glandulosa (Blanco) 
Merr., Dept. of Interior, Manila. 37. 1905. = Thornbera 
dalea (L.) Rydb., Fl. N. Am. 24(2) : 120. 1920. = Dalea 
annua (Mill.) Kuntze, Rev. Gen. 178. 1891. 

Amorpha glauca Raf., Aut. Bot. 80. 1840. [Missouri.] A 
synonym of A. fruticosa L. s.lat. as suggested by Merrill 
(Index Rafinesquianus, p. 142. 1949.) 



i<>75] Amorpha — Wilbur 40/ 



Amorpha Lewisii Loddiges ex Loudon, Arb. Brit. 2: 607. 



1888. [Public 
in synonymy.] 



portedly 



Amorpha ludoviciana Hort. ex Lavallee, Arb. Segrez. 60. 

1877. [Publication not seen by me but this binomial is 

reportedly a nom. nud.] 
Amorpha Ludwigii Hort. ex K. Koch, Dendr. 1: 70. 1869. 

[Publication not seen by me; reportedly published in 

synonymy.] 
Amorpha ? lutea Raf., Fl. Ludov. 105. 1817. [Louisiana.] 

Clearly not an Amorpha but its identity has not yet been 

established. 

Amorpha macrophylla Raf., Aut. Bot. 79. 1840. [Florida.] 
Probably another .synonym of A. fruticosa L. s.lat. as 
suggested by Merrill (Index Rafinesquianus, p. 142. 
1949.). The identity of this name is totally impossible to 
surmise if one were following Palmer's treatment with 
its numerous accepted segregates from the Amorpha 
fruticosa complex. 

Amorpha marginata Hort. ex Lavallee, Arb. Segrez. 60. 
1877. [Publication not seen by me but reportedly the 
binomial appeared as a nom. nud.] 

Amorpha non-pcrforata Schkuhr, Bot. Handb. Deutschl. 
Gew. 2: 333. 1796. [Publication not seen. Usually listed 
in the synonymy of A. fruticosa L.] 

Amorpha ornata Wenderoth, Ind. Sem. Hort. Marburg. 
1835. [Publication not seen by me; reportedly a nom. 
nud.] 

Amorpha oralis M. E. Jones, Contr. West. Bot. no. 16: 
32. 1930. Typification : Arizona: Miller Canyon, Hua- 
chuca Mts., M. E. Jones 25027 (holotype, not seen; iso- 
types, cas!, mo!, okla!, rm !, uc!) = Indigofera sphae- 
rocarpa A. Gray. 

Amorpha pedalis Blanco, Flora de Filipinas, ed. 1. 553. 
1837. [Philippines.] According to Merrill (Dept. of In- 
terior, Manila. 19. 1905) this is probably Solominia 
oblongifolia DC. (Polygalaceae). 



408 



Rhodora [Vol. 77 



/ 



333. 1796. [Publication not -seen. Usually listed in the 
synonymy of A. fruticosa L.] 

Amorpha rabiae Lexarza, Nov. Veg. Desc. fasc. I. 22. 1824. 
[Michoacan, Mexico.] Palmer (Jour. Arn. Arb. 12: 197. 



1931) presents a resume 



It 



does not possess the characters of an Amorpha but its 
identity is otherwise uncertain. 

Amorpha return Raf., Aut. Bot. 80. 1840. [Appalachian 
Mts.] Probably a .snyonym of A. fruticosa L. ; Rafines- 
que's description mentioning its tomentose leaflets and 
acute calyx-lobes would certainly exclude the possibility 
of its being A. glabra as was suggested by Merrill (Index 
Rafinesquianus, p. 142. 1949) and Rehder (Bibliogr. 
Cult. Trees and Shrubs. 369. 1949). 

Amorpha tomentosa ? Raf., Fl. Ludov. 105. 1817. [Louisi- 
ana.] Identity uncertain as all that Rafinesque states 
about it is "a doubtful species, of which Robin only men- 
tions it being tomentose . . ." Rydberg (N. Am. Fl. 24 
(1) : 27. 1919) questioningly placed it in the synonymy of 
A. paniculata but the description is far too incomplete 
to allow anyone to identify the plant being described. It 
is not even certain that it is an Amorpha. 

Amorpha tomentosa Raf., Aut. Bot. 81. 1840. not Raf., Fl. 
Ludov. 105. 1817. ["On Wabash and West Kentucky."] 
Merrill (Index Rafinesquianus, p. 142. 1949) question- 
ingly suggests that it might be a synonym of A. canes- 
reus Pursh but that species is unknown to me from 
"West Kentucky" one of the two cited localities. 

Amorpha verrucosa Raf. Aut. Bot. 80. 1840. [Alabama.] 
Merrill (Index Rafinesquianus, p. 142. 1949) suggested 
this binomial was a synonym of A. fruticosa L. and I 

would agree that the original description is vague enough 
to fit the very broad concept I have of the species, al- 
though it would be impossible to tell to which of the 
segregates recognized by Palmer from the A. fruticosa 
complex that Rafinesque's species belongs. 



1975] Amorpha — Wilbur 409 



LITERATURE CITED 



ABRAMS, L. 1944. Illustrated Flora of the Pacific States. Amorpha 



ford 



MUNZ, P. A. 1959. A California Flora. Amorpha on p. 852. Univ. 



of California Press. 



Palmer, E. J. 1931. Conspectus of the Genus Amorpha. Jour. 



Arnold Arb. 12: 157-197. 



Rydberg, P. A. 1919. [Treatment of Amorpha] in the N. Amer. 



PL 24: 2G-34. 



Schneider, C. K. 1907. Conspectus Generis Amorphae. Bot. Gaz. 



■13: 297-307. 



DEPARTMENT OF BOTANY 
DUKE UNIVERSITY 
DTRHAM, N.C. 2770G 



PAPAVER IN NORTH AMERICA 

NORTH OF MEXICO 1 



Robert W. Kiger- 



com 



The following diagnostic treatment includes the native 
and introduced species of Papaver found in North America 
north of Mexico. It is primarily synthetic in nature and 
is based in large part on the works cited under "Litera- 
ture," as well as on the collection of specimens in the 
United States National Herbarium, Smithsonian Institu- 
tion. This treatment is not offered as a definitive revision, 
the achievement of which will require extensive study in 
the field, herbarium and laboratory. In some cases, exist- 
ing collections are very inadequate, and this is especially 
true of the problematic scapose poppies. In view of the 
present confused situation with regard to the latter in 
North America, a broad species concept is here adopted 
for them. In this arctic and alpine 

of dubious merit have been proposed, some on the basis 
of only one or a few specimens and with little regard for 
the overall generic context. Much importance has been 

numbers in some instances, but until 
al patterns in Papaver are fully in- 
vestigated and their relationships to morphological patterns 
analyzed, such data contribute little to sound taxonomic 
circumscriptions and dispositions. 

When the native North American species of Papaver 
are better known, it is probable that some taxa herein 
submerged in broadly conceived species will be recognized 
separately at the specific and varietal levels. Until such 
time, however, practicality and fidelity to what is actually 

a Based on a study conducted for the Flora North America Program, 
supported by National Science Foundation Contract C-757 with the 
Smithsonian Institution. 

-Present address: Hunt Institute for Botanical Documentation, 
Carnegie Mellon University, Pittsburgh, Pennsylvania 15213. 



omosome 



com 



410 



1975] Papaver — Kiger 411 



known about the native poppies seem best served by a 
synoptic approach utilizing these broad taxa. I have made 
no attempt to relate the introduced poppies to the many 
infraspecific taxa that have been proposed for those species 
in their native ranges. This would be virtually impossible 
on the basis of the morphologies of most individual plants 
found in North America. Any such correlations which 
could be made would be essentially meaningless in the 
North American context, within which, due to geographic- 
ally random introductions and subsequent hybridizations, 
there is no biologically significant pattern of variation in 
evidence. 

The sectional nomenclature utilized herein follows that 
presented in my recent review of the subject (Kiger, 1973). 
The circumscriptions and descriptions of sections generally 
follow the traditional ones of existing revisions, particu- 
larly that by Fedde (1909). Only characters of species 
found in North America north of Mexico are included in 
the sectional descriptions. The descriptions of all taxa 
follow the outline and glossary developed for Flora North 
America (Porter, et ah, 1973). 



Papaver L., Sp. PL, 506. 1753; Gen. PI. ed. 5, 224. 1754. 

Plants annual or perennial, herbaceous, caulescent, sub- 
scapose or scapose, monoclinous, sap gummy, opaque. Main 
stems eramous or ramose. Leaves alternate, sometimes 
basally rosulate; blades entire to bipinnatipartite. Flowers 
solitary, pedicellate or scapose, actinomorphic, complete, 
syncarpous, apostemonous. Sepals 2, free, caducous, entire. 
Petals 4, entire, aestivation corrugate. Stamens numerous. 
Carpels 3-18; styles absent; stigmas 3-18, sessile, radiate 



mem 



often 



conspicuous. Fruits capsular, dehiscence poricidal, pores 
subapical, 3-18, locules 3-18. 



412 



Rhodora [Vol. 77 



KEY TO SPECIES 



a. Plants caulescent or rarely subscapose, at least a few 
cauline leaves present, b. 



■ ■ 



b. Upper cauline leaves amplexicaulous 



1. P. somniferum. 

b. Upper cauline leaves not amplexicaulous. c. 

c. Ovaries and capsules setose. . . 5. P. hybridum. 
c. Ovaries and capsules glabrous, d. 

d. Plants perennial ; stems eramous ; petals 5 cm 

or more long 6. P. orient ale. 

d. Plants annual ; stems ramose ; petals less than 
5 cm long. e. 
e. Stigmatic disc depressed-conic, usually 

umbonate 4. P. calif ornicum. 

e. Stigmatic disc essentially plane, f. 

f. Pedicels markedly patent hispid dis- 
tally; capsules less than twice as long 

as broad 2. P. rhoeas. 

f. Pedicels strongly appressed hispid 
distally; capsules twice as long as 
broad 3. P. dubium. 

a. Plants scapose, leaves all basal, g. 

g. Leaves entire to 3 (rarely 5)-lobed, essentially gla- 
brous ; capsules obovoid-obconic. ... 9. P. ivalpolei. 

g. Leaves pinnatipartite or bipinnatipartite with more 
than 5 lobes, variously vestite, at least when young; 
capsules narrowly clavate to globose, h. 

h. Setae on ovaries and capsules basally tubercu- 
late. i. 
i. Petals salmon pink to yellow, paler basally; 

Rocky Mountains 7. P. alpinum. 

i. Petals white to rose with yellow basal spot; 

Alaska 8. P. alboroseum. 

h. Setae on ovaries and capsules not basally tuber- 
culate. j. 

j. Stigmatic disc distinctly conic and/or um- 
bonate. k. 



1975] Papaver — Kiger 413 



k. Leaves mostly bipirmatipartite; capsule 
obovoid 11. -P. mcconnellii. 

k. Leaves mostly pinnatipartite ; capsule nar- 
rowly oblong-cylindric to clavate 

10. P. macounii. 



j. Stigmatic disc essentially plane. 1. 

I. Mature plants over 25 cm tall; petals 4-6 
cm long; scapes glabrate to sparsely ves- 
tite. 13. P. nvdicaule. 

1. Mature plants less than 25 cm tall; petals 
to 4 (rarely to 5) cm long; scapes moder- 



ately to densely vestite 



• • 



12. P. lapponicum ssp. occidental?. 



Sect. Papaver 



Plants annual, caulescent, glaucous, glabrate. Leaf 
blades simple or pinnatipartite. Basal leaves petiolate. Cau- 
line leaves sessile, upper amplexicaulous. Filaments cla- 
vate. Stigmatic disc essentially plane. Fruits globose or 
subglobose, glaucous, glabrous. 



1. P. somniferum L., Sp. PL, 508. 1753. 

Plants 3-15 dm tall. Main stems distally cauliramous, 
stout; branches few. Leaf blades simple and coarsely bi- 
dentate, or pinnatipartite with lobes irregularly dentate. 
Pedicels glabrate or sparsely pale setose distally. Petals 
white, pink, red or purple, darker spotted basal ly, 3.5-6.0 
cm long. Filaments white; anthers pale yellow. Stigmas 
5-18. disc margin deeply lobed, membrane conspicuous. 
Fruits usually stipitate, 3-0 cm long, not ribbed. 

Adventive locally throughout the United States and 
Canada as far north as Newfoundland, introduced from 
Europe, native of Eurasia; fields, roadsides and waste 
places, especially about habitations ; flowering June-Septem- 
ber. 



414 Rhodora [Vol. 77 



Sect. Rhoeades Bernh. 



Plants annual, caulescent or subscapose, glabrous, pilose, 
hirsute, hispid or setose. Leaf blades pinnatipartite or 
bipinnatipartite. Basal leaves petiolate. Cauline leaves ses- 
sile, not amplexicaulous. Filaments filiform. Stigmatic disc 
essentially plane, sometimes umbonate, margin lobed, lobes 
rounded to obtuse. Fruits glabrous. 



2. P. rhoeas L., Sp. PL, 507. 1753. 

Plants caulescent, hispid or setulose, 2.5-9.0 dm tall. Main 
stems distally cauliramous, slender to stout. Leaf blades 
pinnatipartite or bipinnatipartite, lobes acuminate, coarsely 
dentate. Cauline leaves often somewhat clustered beneath 
the pedicels. Pedicels sparsely to moderately patent hispid 
or setulose. Petals pink to red, sometimes darker spotted 
basally, 2.0-4.5 cm long. Filaments purple; anthers bluish. 
Stigmas 5-18, disc essentially plane, broader than capsule 
apex, membrane conspicuous. Fruits slightly stipitate or 
sessile, broadly obovoid or subglobose, 1-2 cm long, ob- 
scurely ribbed, glaucous. 

Adventive locally throughout the United States and 

Canada as far north as Alaska, introduced from Europe; 

fields, roadsides and disturbed sites, especially about habi- 
tations; flowering March-October. 



3. P. dubium L.. Sp. PL, 1196. 1753. 

Plants caulescent or subscapose, 2-6 dm tall. Main stems 
basicauliramous, hirsute to hispid. Basal leaf blades pin- 
natipartite, glaucescent, hispid. Cauline leaf blades bipin- 
natipartite, glaucescent, hispid. Pedicels distally appressed 

hispid, proximally patent hispid. Petals red, sometimes 
darker spotted basally. Filaments purple; anthers violet. 
Stigmas 7-9, disc essentially plane. Fruits sessile or slightly 
stipitate, obovoid, L:W::2:1, usually distinctly ribbed, 
glaucous. 



1 975] Papaver — Kiger 415 



Naturalized locally throughout the contiguous United 
States, introduced from Europe; fields, roadsides, thickets, 
disturbed sites, especially near habitations; flowering May- 

Aufirust. 



4. P. californicum A. Gray, Proc. Amer. Acad. 22: 313. 
1887. 

P. lew man ii Greene 

Plants caulescent, glabrous or sparsely pilose, 3-6 dm 
tall. Main stems distally cauliramous, slender. Leaf blades 
pinnatipartite or bipinnatipartite, lobes sometimes dentate. 
Petals red, greenish spotted basally, 1-2 cm long. Fila- 
ments greenish yellow; anthers yellow. Stigmas 5-8, disc 
plane to depressed-conic, usually umbonate, not broader 
than capsule apex, membrane conspicuous. Fruits sessile, 
ellipsoid to obovoid-turbinate, 1-2 cm long. 

Native, endemic to western California; chaparral and 
oak woodlands in the mountains at elevations below 800 m, 
especially in clearings, burns and other disturbed sites; 
flowering April-May. 



Sect. Argemonidium Spach 

Plants annual, caulescent, pilose, villous or hispid. Leaf 
blades pinnatipartite or bipinnatipartite. Basal leaves pe- 
tiolate. Cauline leaves sessile, not amplexicaulous. Fila- 
ments clavate. Stigmatic disc convex and vaulted, lobes 
obtuse. Fruits setose. 



5. P. hybridum L., Sp. PL, 506. 1753. 

P. apulum Ten. var. micranthum (Boreau) Fedde 

misappl. 

Plants 1-5 dm tall. Main stems basiramous and cauli- 
ramous. Pedicels moderately to densely appressed to 
spreading hispid. Petals red, darker spotted basally. Fila- 
ments dark violet: anthers pale blue. Stigmas 4-8, disc 
deciduous at maturity. Fruits sessile, obovoid to subglobose, 



416 



Rhodora [Vol. 77 



1.0-1.5 cm long, obscurely to distinctly ribbed, vestiture 
recurved-arcuate, spreading or ascending. 

Naturalized locally in California, introduced from Eura- 
sia; fields, vineyards and disturbed sites; flowering March- 
May. 

Specimens attributed to P. apulum var. micranthum in 
herbaria and reported as such in Munz and Keck (1959) 
and in Munz (1968) belong here. 



SECT. Oxytona Bernh. 

Plants perennial, robust, caulescent, hispid. Leaf blades 
pinnatipartite. Basal leaves petiolate. Cauline leaves ses- 
sile, not amplexicaulous. Flowers very large. Filaments 
clavate. Stigmatic disc plane. Fruits subglobose, glaucous, 
glabrous. 



6. P. orientale L., Sp. PL, 508. 1753. 

Plants 6-10 dm tall. Main stems eramous. Leaf lobes 
aristate. Pedicels moderately to densely appressed pale 



hispid. 



sometime 



5-8 cm long. Filaments purple; anthers violet. Stigmas 



13-18. 



cm 



Adventive locally in the middle-Atlantic United States, 
escaping from cultivation, introduced from Europe, native 
of southwest Asia; fields and disturbed sites; flowering in 
May. 

Papaver bracteatum Lindl., which some consider distinct 

from P. orientale, has deep red petals and has foliaceous 
bracts subtending the sepals. It is occasionally cultivated 
and might be found as an escape. 



Sect. Lasiotrachyphylla Bernh. 

Plants perennial, scapose. Main stems eramous. Basal 
leaves petiolate, rosulate, blades simple, pinnatipartite or 
bipinnatipartite. Cauline leaves absent. Stigmatic disc 
plane to conic, sometimes vaulted, sometimes umbonate. 
Fruits setose. 



1975] 



Papaver — Kiger 417 



7. P. alpinum L., Sp. PL. 507. 1753. 

P. pygmaeum Rydb. 

P. nudicaule L. ssp. radicatitm (Rottb.) Fedde var. 
pseudoeorydali folium Fedde 

Plants cespitose. Leaf blades pinnatipartite or bipin- 
natipartite, glabrate or sparsely hispid. Scapes erect, to 

10 cm tall, sparsely hirsute. Petals yellow to salmon pink, 
paler spotted basally, to 1 cm long. Stigmas usually 5, disc 
vaulted. Fruits obovoid-cylindric or ellipsoid, whitish 
setose, trichomes basally tuberculate. 



M 



om 



ing July-August. 

For discussion of the relationship of these plants to 
European and Asian members of the complex see D. Love 

(1969). 

8. P. alhoroseum Hult, Fl. Kamtchatka 2, Sv. Vet.-Akad. 
Handl. ser. 3, 5, no. 2: 141, t. 3, f. c. 1928. 

Plants cespitose, caudex short. Leaf blades bipinnati- 
partite. pale setose above and beneath, primary lobes 2-5- 
partite. Scapes arcuate-ascending, to 15 cm tall, setose. 
Petals white to rose, yellow spotted basally. Stigmas 5-6, 
disc essentially plane. Fruits ovoid to globose, ribbed, 
whitish to brown setose, trichomes basally tuberculate. 

Native to the Kenai Peninsula, Alaska, also to Kamt- 
chatka, U. S. S. R. ; sandy and gravelly soils at low eleva- 
tions. 



9. P. walpolei Pors., Rhodora 41: 231. 1939. 

Plants densely cespitose. Leaf blades entire or pinnati- 
partite. subcoriaceous, glabrous, lobes (when present) 3, 
rarely 5, broadly obtuse, margins revolute, petiole bases 
conspicuously persistent. Scapes erect, to 16 cm tall, hir- 
sute or hirtellous distally. Petals pale yellow or c: 



earn 



418 Rhodora [Vol. 77 



yellow spotted basally, to 2.5 cm long. Stigmas usually 5, 
disc conic, broader than capsule. Fruits obovoid-obconic, 
1.5 cm long, pale yellowish setose or setulose, trichomes 
slender, basally tuberculate, sometimes only obscurely so. 
Native to the Seward Peninsula, Alaska, also to north- 



moun 



tain slopes in gravelly loam and solifluction soils, often of 
limestone origin; flowering May-August. 



10. P. macounii Greene, Pittonia 3: 247. 1897. 

P. keelei Pors. 

P. alaskanum Hult. var. macra?ithum Hult. 

P. scammianum D. Love 

P. hultenii Knaben 

P. microcarpum DC., misappl. 

P. nudicaule L., misappl. 

Plants solitary to densely cespitose. Leaf blades pinnati- 
partite or rarely bipinnatipartite, glabrate to pilose. Scapes 
erect, to 4 dm tall, moderately pilose. Petals yellow. 
Stigmas 3-5, disc conic, umbonate, sometimes vaulted. 
Fruits narrowly oblong or clavate, L:W::2:1, setose, tri- 
chomes not basally tuberculate. 

Native and widespread in Alaska, Yukon Terr., North- 
west Terr, and British Columbia ; sandy and gravelly soils, 
heaths, thickets, meadows, often on slopes, to over 2100 m. 



11. P. mcconnellii Hult., Fl. Alaska & Yukon 5, Lunds Univ. 

Arssk. N. F. Avd. 2, 41, no. 1 : 803, /. 1. 1945. 

Plants cespitose, to 15 cm tall. Leaf blades bipinnati- 
partite, rarely pinnatipartite, glaucous, sparsely pilose 
above and beneath. Scapes erect, pilose. Petals yellow. 
Stamens usually shorter than mature ovary. Stigmatic 
disc convex, distinctly apiculate-umbonate, membrane con- 
spicuous. Fruits obovoid, pale setose, trichomes not basally 
tuberculate. 

Endemic to Yukon Terr. ; sandy and gravelly soils. 



1975] 



Papaver — Kiger 419 



12. P. lapponicum (Tolm.) Nordh. ssp. occidentale (Lundstr.) 

Knaben. Op. Bot. 2, 3: 55. 1959. 

P. radicatum Rottb. ssp. lapponicum Tolm. 

P. radicatum ssp. occidentale Lundstr. 

P. lapponicum ssp. porsildii Knaben 

P. nudicaule L. ssp. radicatum (Rottb.) Fedde var. 

coloradense Fedde 
P. nudicaule ssp. radicatum var. columbianum Fedde 

P. kluanensis D. Love 

P. freedmanianum D. Love 

P. nigroflarum D. Love 

P. cormuallisensis D. Love 

P. alaskanum Hult. var. alaskanum 

P. deualii G.iaerevoll 

P. radicatum Rottb. ssp. radicatum, misappl. 

Plants cespi'tose. Caudex short to elongate. Leaf blades 
mostly bipinnatipartite, pilose; petiole bases usually per- 
sistent, sometimes conspicuously so, pale to dark brown. 
Scopes ascending to erect, to 20 (rarely 25) cm tall, sparse- 

i n vn+ n i,r nnnro^cH fr> r^tpnt Vn'snid or nilose. Petals 



mo 



yellow or rarely white. Stigmas 5-8, disc plane to slightly 
convex, sometimes vaulted. Fruits subglobose, ellipsoid or 
obovoid, setose, trichomes not basally tuberculate. 

Native and widespread in arctic North America from 
Greenland to Alaska, extending southward in disjunct 
populations in the Rocky Mountains to Colorado and north- 
ern New Mexico at high elevations, also in Lapland; sandy 
and gravelly soils, often on talus, arctic and alpine tundra 

to ca. 3000 m. 

For discussions of the proper typification of P. radica- 
tum Rottb. and its bearing on the nomenclature in this 
complex see A. Love (1962a, 1962b), Knaben (1958) and 
Knaben and Hylander (1970). The latter interpretation is 
here accepted. See D. Love (1969) for discussion of the 
Rocky Mountain plants of this complex, held separate by 

her. 



420 Rhodora [Vol. 77 



13. P. nudicaule L., Sp. PL, 507. 1753. 

Plants cespitose. Leaf blades pinnatipartite, less fre- 
quently bipinnatipartite, glabrate or setose. Scapes erect, 

25-50 cm tall, stout, glabrate or sparsely hispid. Petals 
yellow, red or white. Stigmas 4-6, disc plane, membrane 
obsolescent. Fruits clavate or obovoid, setose, trichomes 
not basally tuberculate. 

Adventive locally in Alaska and Yukon Terr., 
from cultivation, native to arctic and alpine Asia; road° 
sides and disturbed sites, especially about habitations. 



escaping 



LITERATURE 



Abrams, L. 1944. Illustrated flora of the Pacific States, Washing- 
ton, Oregon and California. Vol. 2. Stanford. 

Anderson, J. P. 1959. Flora of Alaska and adjacent parts of 
Canada. Ames. 

Bocher, T. W., et al. 1906. Gronlands flora, ed. 2. Copenhagen. 
Ernst, W. R. 1962. The genera of Papaveraceae and Fumaria- 

ceae in the southeastern United States. Jour. Arnold Arb 43- 

3 15-343. 

Faberge, A. C. 1944. Genetics of the Scapiflora section of Papaver. 
Ill, Interspecific hybrids and genetic homology. Jour. Genet. 

46: 125-149. 

Fedde, F. 1909. Papaver in Engler, A., ed., das Pflanzenreich 
40(4, 104): 288-386. 

FERNALD, M. L. 1950. Gray's manual of botany, ed. 8. New York. 
Gleason, H. A. 1963. The new Britton and Brown illustrated flora 

of the northeastern United States and adjacent Canada. Vol. 2. 
New York. 

Hulten, E. 1945. Flora of Alaska and Yukon, 5. Lunds Univ. 
Arssk. N. F. Avd. 2, 41, no. 1. 

. 1968a. Comments on the Flora of Alaska and Yukon. 



Ark. Bot. 7. 1. 



• 1968b. Flora of Alaska and neighboring territories. A 
manual of the vascular plants. Stanford. 

Jepson, W. L. 1922. A flora of California. Vol. 1, Part 7. Berke- 
ley. 

Kiger, R. W. 1973. Sectional nomenclature in Pajxiver L. Taxoii 
22: 579-582. 



1975] Papaver — Kiger 421 



Knaben, G. 1958, Papaver-studier, med et forsvar for P. radi- 
catum Rottb. som en islandsk-skandinavisk art. Rlyttia 16: 61- 



80. 



. 1959a. On the evolution of the radi catum-gr ou p of the 

Scapiflora papavers as studied in 70 and 56 chromosome species. 

Part A. Cytotaxonomical aspects. 0]). Rot. 2, 3. 

. 1959b. On the evolution of the radicatum-gr oup of the 



Scapiflora papavers as studied in 70 and 56 chromosome species. 

Part R. Experimental studies. 0]). Rot. 3, 3. 

, & N. Hylander. 1970. On the typification of Papaver 

radicatum Rottb. and its nomenclatural consequences. Rot. Not. 
123: 338-345. 

LOVE, A. 1962a. Typification of Papaver radicatum — a nomen- 
clatural detective story. Rot. Not. 115: 113-136. 

. 1962b. Nomenclature of North Atlantic Papavers. 

Taxon 11: 182-138. 

Love, P. 1969. Papaver at high altitudes in the Rocky Mountains. 
Rrittonia 21: 1-10. 

, & N. J. Freedman. 1956. A plant collection from SW 



Yukon. Rot. Not. 109: 153-211. 
Mowat, A. R., & S. M. Walters. 1964. Papaver in Tutin, T. G., 
et AL., eds., Flora Europaea. Vol. 1, 247-250. Cambridge. 

MUNZ, P. A. 1968. Supplement to a California flora. Rerkeley. 

— , in collaboration with P. P. Keck. 1959. A California 



flora. Rerkeley. 

Polunin, N. 1940. Botany of the Canadian eastern Arctic. Part I. 
Pteridophyta and Spermatophyta. Nat. Mus. Canada Bull. 92. 

. 1959. Circumpolar arctic flora. Oxford. 

POPOV, M. G. 1937. Papaver in KOMAROV, V. L., ED., Flora S. S. 
S. R. Vol. 7, 598-646. Moscow. 

PORSILD, A. E. 1939. Contributions to the flora of Alaska (con- 
tinued). Rhodora 41: 199-254. 

. 1951. Rotany of southeastern Yukon adjacent to the 



Canol Road. Nat. Mus. Canada Bull. 121. 

. 1955. The vascular slants of the western Canadian 



Arctic Archipelago. Nat. Mus. Canada Bull. 135. 

. 1964. Illustrated flora of the Canadian Arctic Archi- 



pelago. Nat. Mus. Canada Bull. 146. 

Porter, D. M., et al. 1973. A guide for contributors to Flora 
North America. Part II. An outline and glossary of terms for 
morphological and habitat description (provisional edition). FNA 
Report GC). Washington. 



422 Rhodora [Vol. 77 



Rydberg, P. A, 1922. Flora of the Rocky Mountains and adjacent 
plains, ed. 2. New York. 

Small, J. K. 1933. Manual of the southeastern flora. New York. 
Wiggins, I. L., & J. H. Thomas. 1962. A flora of the Alaskan 

arctic slope. Arctic Inst. N. Amer. Special Publ. 4. 



DEPARTMENT OF BOTANY 
SMITHSONIAN INSTITUTION 
WASHINGTON, D.C. 20560 



THE NORTH AMERICAN SPECIES, 
BOERHAVIA SPICATA (NYCTAGINACEAE) , 

IN NORTHWESTERN ARGENTINA 



Philip D. Cantino 



Boerhavia spicata Choisy is a morphologically variable 
herb widespread in the Sonoran and Chihuahuan deserts 
of the southwestern United States and northern Mexico, 
and heretofore known only from that region. During the 
course of field study as a worker in the IBP Structure of 
Ecosystems Program, I found an isolated population of 
this species in Catamarca Province, Argentina. This con- 
stitutes the first known occurrence of the species in South 
America, and is an addition to the already sizable list of 
species known to have range disjunctions between the 
desert regions of southwestern North America and north- 
ern Argentina and Chile (Bray, 1900; Johnston, 1940; 
Raven. 1963: Solbrig, 1972). 

* 

Plants of Boerhavia spicata occur abundantly on a small 
group of sand dunes located 35 km west of the town of 
Andalgala, at the north edge of the Bolson de Pipanaco. 
In spite of extensive field work in the Andalgala region 
over an eight-month period in 1972-73, no other popula- 
tions, or even isolated individuals, of the plant were found. 
Several Argentine botanists familiar with the flora of the 
region were consulted, and none had seen the species pre- 
viouslv. In addition, a literature search revealed no refer- 
ences to the occurrence of B. spicata or other species with 
similar characteristics in South America. 



I considered the possibility that the population 



m 



represent an undescribed species. An inspection of the 
worldwide collection of Boerhavia at the Harvard Univer- 
sity Herbaria revealed a strong resemblance between the 
Argentine specimens and members of the B. spicata 
complex (explained below). No marked resemblance was 
found to any other species of Boerhavia. 



423 



424 Rhodora [Vol. 77 



The Boerhavia spicata complex has been considered by 



om 



a number of authors (Standley, 1909; Tidest: 
tell, 1941; Kearney and Peebles, 1960; Ferris, 1964) to 
consist of three to four distinct species : B. spicata Choisy, 
B. watsonii Standley, B. coulteri (Hook, f.) S. Watson, and 
B. torreijana (S. Watson) Standley. More recently it has 
been interpreted as a single morphologically variable 
species, B. spicata Choisy (Reed, 1969). Upon examina- 
tion of the Harvard Herbaria collection of the complex 
(50 specimens), I tend to agree with Reed's interpretation 
that we are dealing with a single variable species. The 
characters used by the above-mentioned authors to dis- 
tinguish the four species (e.g., degree of crowding of the 
flowers in the inflorescence; shape of the ridges and furrows 
of the anthocarp; degree of rugosity of the anthocarp; 
presence or absence of glandular pubescence on the stems 
and glandular dots on the foliage) all show gradual varia- 
tion between the extreme character states. However, the 
species should, perhaps, be segregated into two varieties 
— one, with a center of distribution in the Sonoran Desert, 
having a tendency towards closely spaced flowers, acute 
to acuminate leaf-apices, and densely glandular-villous 
stems; and the other, with a center of distribution in the 
Chihuahuan Desert, having a tendency towards loosely 
spaced flowers, obtuse leaf apices, and puberulent stems 
with few or no glands. The Argentine population clearly 
has affinities with the latter variety. 

The above analysis is intended as a suggestion only. I 
do not feel justified in dividing the species into varieties 
based solely on the very limited herbarium study that I 
have done. Crossing experiments in the laboratory and a 
field study of ecotypic variation would help greatly to 
clarify the taxonomic relationships within this difficult 

)lex. Meanwhile, for the purposes of identifying the 
Argentine population, I am following Reed in his place- 
ment of the entire complex in the species Boerhavia spicata 
Choisy. The Argentine plants are clearly members of the 
complex; thus until someone does a definitive revision of 
the genus, they should be placed in B. spicata Choisy. 



com 



p: 



1975] Boerhavia — Cantino 425 



The following description is patterned after Reed's de- 
scription of Boerhavia spicata but has been modified so as 
to describe the Argentine population rather than the 
species as a whole: 



Erect or decumbent annual from a slender taproot, usu- 
ally with many branches spreading from the base; stems 
2-6 dm long, much branched, often tinged with pink, white- 
puberulent on the lower parts (occasionally sparsely glan- 
dular-hirtellous at the nodes only), glabrous on the upper 
parts; leaves opposite, the petioles puberulent, 0.2-3.0 cm 
long, the leaf-blades oval to ovate or ovate-lanceolate, 0.8- 
4.0 cm long, 0.4-2.5 cm wide, truncate to obtuse or slightly 
oblique at base, obtuse to acuminate at the apex, the margins 
entire to sinuate, green above, paler beneath, brown-punc- 
tate or not, sparsely puberulent to glabrate; inflorescence 
cymose-paniculate, spreadingly much-branched, the branches 
slender, glabrous; flowers on short pedicels 0.3 to 1.0 mm 
long, loosely spaced on the ultimate branches of the inflores- 



m 



mm 



stamens 2, short-exserted : anthocarp narrowly obovoid, 
3 mm long, 1 mm wide, rounded at apex, acute at the 
base, 5-angulate, the sulci nearly closed, strongly rugose. 

Specimens (Cantino fi64, 724) are deposited in the her- 
baria of the following institutions in the United States 
and Argentina : Harvard University, University of Michi- 
gan, University of Arizona, New Mexico State University, 
University of Texas, Instituto Miguel Lillo (Tucuman), 
Universidad Nacional de Cordoba, and Instituto de Botan- 
ica Darwinion (San Isidro). 



ACKNOWLEDGMENTS 



I wish to express my appreciation to Dr. Otto T. Solbrig, 
Dr. Reed C. Rollins, Dr. Carroll E. Wood, and Dr. Rolla 
M. Tryon for their advice and encouragement in the prepa- 
ration of the manuscript. 



426 Rhodora [Vol. 77 



LITERATURE CITED 



Bray, W. L. 1900. The relations of the North American flora to 

that of South America. Science II. 12: 709-716. 
Ferris, R. S. 1964. Nyctaginaceae. Pp. 476-482. In F. Shreve & 

I. L. Wiggins. Vegetation and flora of the Sonoran Desert. VoL 1. 



Stanford University Press, Stanford, Calif. 



JOHNSTON, I. M. 1940. The floristic significance of shrubs common 
to North and South American deserts. Jour. Arnold Arb. 21: 
356-363. 

Kearney, T. H., & R. H. Peebles. 1960. Arizona Flora. Univ. of 

Calif. Press, Berkeley & Los Angeles. {Boerhavia, pp. 275-277) 
Raven, P. H. 1963. Amphitropical relationships in the floras of 

North and South America. Quart. Rev. Biol. 38: 151-177. 
Reed, C. F. 1969. Nyctaginaceae. Pp. 203-220. In C. L. Lundell, 

Flora of Texas. Vol. 2. Texas Research Foundation, Renner, 

Texas. 

Solbrig, O. T. 1972. New approaches to the study of disjunctions 
with special emphasis on the American amphitropical desert dis- 
junctions. Pp. 85-100. In P. Valentine, ed. Taxonomy, phyto- 
geography, and evolution. London. 

. 1972. The floristic disjunctions between the "Monte" 

in Argentina and the "Sonoran Desert" in Mexico and the United 
States. Ann. Missouri Bot. Card. 59: 218-223. 



Standley, P. C. 1909. The Allioniaceae of the United States with 
notes on Mexican species. Contr. U.S. Natl. Herb. 12: 303-389. 

Tidestrom, I., & T. Kittell. 1941. A flora of Arizona and New 
Mexico. Catholic University of America Press, Washington, 
D.C. (Boerhavia, pp. 652-655) 



GRAY HERBARIUM 

HARVARD UNIVERSITY 

CAMBRIDGE, MASSACHUSETTS 02138 



GEOGRAPHICAL DISTRIBUTION OF THE 
CEDAR GLADE ENDEMIC VIOLA EGGLESTONII 



Jerry M. Baskin and Carol C. Baskin 



Viola egglestonii Brainerd is a perennial, stemless, blue 
violet that heretofore was thought to be endemic to the 
cedar glades of central Tennessee. Distribution maps pub- 
lished by Russell (1958, 1965) show the species as occurring 
in four counties, and Baskin and Baskin (1970) list five 
additional counties making a total of nine in the Central 
Basin of Tennessee. These counties include Bedford, 
Cannon, Davidson, Marshall, Maury, Rutherford, Smith, 
Sumner and Wilson. Viola egglestonii previously has been 
reported from Warren (Brainerd, 1921) and Hart (Braun, 
1943) counties in south-central Kentucky, but Russell 
(1965) in his extensive treatment of the violets of central 
and eastern United States could not find specimens to 
verify its occurrence in Kentucky. The purpose of this 
note is to report the discovery of populations of V. eggle- 
stonii in cedar (limestone) glades in Kentucky, Georgia 

and Alabama. 

On 15 April 1972 we discovered a population of Viola 

egglestonii in a cedar glade area in Bullitt Co., Kentucky, 
along Co. Rd. 1604, 0.3 miles south of St. Rd. 480 (Baskin 
& Raskin 1191). Two additional populations were found 
on 23 April 1972 in cedar glades in Bullitt Co. The loca- 
tions of these two populations are: Woodsdale Rd., 1.3 
miles east of Deatsville Rd. (Baskin & Baskin 1225) ; 
Bells Mill Rd., 2.6 miles east of U.S. 61 (Baskin & Baskin 
1234). An additional population was found in Bullitt Co. 
on 1 April 1973 on Clarks Lane, 0.7 miles north of St. Rd. 

480 (Baskin & Baskin 1639). 

Populations of Viola egglestonii were found in cedar 
glade areas in Walker and Catoosa counties, Georgia, on 
26 May 1972. The locations of these populations are as 
follows : Walker Co., along U.S. 27, 5.1 miles south of St. 
Rd. 2 (Baskin & Baskin 1301) ; Walker Co., along U. S. 27, 



427 



128 



Rhodora 



[Vol. 77 




Fig. 1. Geographical distribution of Viola egglestonii. Each dot 
represents a county where the species is known to occur. 



4.8 miles south of St. Rd. 2 (Baskin & Baskin 1306) ; Catoosa 
Co., Chickamauga National Park, 2 miles east of Ft. Ogle- 
thorpe on Co. Rd. 2 (Bcuskin & Baskin 1308). 

On 27 May 1972 a population of Viola egglestonii was 
found in a cedar glade in Franklin Co., Alabama. This 
population occurs along Co. Rd. 83, 0.6 miles north of St. 
Rd. 24 (Baskin & Baskin 1328). 

The present known geographical distribution of Viola 
egglestonii is shown in Fig. 1. Herbarium specimens from 



1975] Viola — Baskin & Baskin 429 



the new locations reported in this note have been sent to 
the herbarium at Vanderbilt University (VDB). 



LITERATURE CITED 



RUSSELL, N. H. 1958. The violets of Tennessee. Castanea 23: 

63-76. 
. 1905. Violets (Viola) of central and eastern United 

States. Sida 2: 1-113. 
Raskin, J. M., & C. C. Raskin. New county records for the cedar 
glade endemic Viola egglentonii. Jour. Tenn. Acad. Sci. 45: 

60-61. 
RRAINERD, E. 1921. Violets of North America. Vt. Agri. Exp. 

Sta. Bull. 224. 
BRAUN, E. L. 1943. An annotated catalogue of spermatophytes of 

Kentucky. 161 pp. Published by the author, Cincinnati, Ohio. 



SCHOOL OF BIOLOGICAL SCIENCES 
UNIVERSITY OF KENTUCKY 
LEXINGTON, KENTUCKY 40506 



RECENT ADDITIONS TO THE MARINE ALGAL 
FLORA OF NAHANT, MASSACHUSETTS 1 

E. E. Webber 

The only comprehensive volume to date treating the 
benthic marine algae of northeastern North America is 
that of Taylor (1957). In addition to the systematics and 
species descriptions of relevant plants, Taylor also presents 
a brief historical summary of marine phycology from 



m 



time, however, studies of benthic mar 



New 



summati 



recent research dealing with algae of the sublittoral, with 
those of estuaries and adjacent salt marshes, and with the 
autecology of several species, has been published (Webber, 
1975). This paper also contains a preliminary tabulation 
of 107 benthic algae specifically from the vicinity of the 
Marine Science Institute at Nahant. 

In the early summer of 1973 nineteen additional species 
were collected, mostly from the Nahant rocky shore, and a 
few from a nearby salt marsh. These are listed below. 

References used in determinations of the species were: 
Farlow (1882), Koster (1955), Taylor (1957), Bliding 
(1963), Cardinal (1964), Wilce (1965), and Drouet (1968). 
Extended comments on these species will be reserved for 
forthcoming publications. 

LIST OF SPECIES 
Cyanophycophyta 

Oscillatoria lutea Ag. — Forming a 2 mm thick mat on 
deeply shaded and moist rocks just above high tide 
mark; mixed with Microcoleus vacfinatus (Vauch.) Gom. 

Rivularia atra Roth — epilithic in 2 small rock pools in 
spray zone. 

Tolypothrix sp. and Ciflindrospermum sp. — collected from 
salt marsh soil, mid-littoral. 



Contribution No. 14 from the Marine Science Institute, Nahant, 



Massachusetts 01908. 



430 



1975] Nahant Algae — Webber 431 



Chlorophycophyta 

Tetraspora sp. — Located at the bases of the marsh grass, 
Spartina patens Ait. (Muhl.) upper littoral. 

Pring sheimiella seutata (Reinke) Schmidt et Petrak — col- 
lected once, attached to the operculum of Littorina 



obtusata. 



ma 



(J. Ag.) Dangeard ssp. marginata 



comm 



B. marginata ssp. subsalsa (Kjell.) Bliding — located im- 
mediately above B. marginata. 

Enterow orpha compressa (L.) Grew var. compressa — 
occurs in quantity in a splash zone pool at the top of 
the very exposed East Point rocks. 

E. ahhieriana Bliding — common in tide pools at high tide 

level. 

E. clathrata (Roth) Grev. (type I)— attached to small 
stones in the lower littoral zone of Lynn Harbor. 

E. clathrata (Roth) Grev. (type II) — entangled with 
CJadophora sp. in the lower littoral zone of Lynn Harbor. 

E. flexuosa (Wulfen ex Roth) J. Ag. ssp. flexuosa — grow- 
ing on shells in the lower littoral zone of Lynn Harbor. 

Urospora coUabcns (C. Ag.) Holmes et Batters — com- 
monly enilithic in the mid-littoral zone at Nahant, often 



m 



Codiolum gregarium A. Braun. 



Rhizoclonium riparium (Roth) Harv. f. riparium — while 
R. tortuosum is the common species of this genus at 
Nahant, R. riparium was located in one splash -zone tide 
pool at the East Point rocks. 



Phaeophycophyta 

Ectocarpus fasciculatus var. refractvs (Kutz.) Ard. — 
common on Layninaria stipes; previously reported by 
Taylor (1957) only from Maine, as an epiphyte on 
Porphyra. 

Myriotrichia filiformis Harv. — epiphytic on Chordaria 
flag ellif ormis in a lower littoral zone tide pool. 



432 " Rhodora [Vol. 77 



Saceorhiza dermatodea (De la Pyl.) J. Ag. — epilithic at 

8 to 20 foot depths, mixed with Laminaria saccharine 
(Wilce, 1965) and L. digitata. Published accounts of 
S. dermatodea along* the northeast coast suggest that this 
species is limited to coastal waters north of Cape Ann, 
Massachusetts (Farlow, 1882; Lamb and Zimmermann, 
1964; Stone, et. al, 1970; Mathieson and Fralick, 1972). 
However, records for the distribution for Saceorhiza in 
New England (Taylor, personal communication) reveal 
that plants of this species extend southward to the Cape 
Cod (Sandwich, Massachusetts) area. Indeed, S. derma- 
todea is known from the sublittoral at both ends of the 
Cape Cod Canal (Wilce, personal communication). 



Rhodophycophyta 

Nemalion multifidum (Weber et Mohr) J. Ag. — epilithic 
in the upper sublittoral, uncommon, only female plants 

located. 



ACKNOWLEDGMENTS 

Appreciation is expressed to Dr. W. Randolph Taylor 
for sending me collection records of Saceorhiza dermatodea 
from his personal herbarium. Plants of this species were 
collected simultaneously at Nahant (July 14, 1973) by 
Jane Menge and Barry Spracklin. 

In addition, Myriotrichia filiformis, Nemalion multifidum, 
and Rivularia atra were collected initially by Jane Menge, 
while Prigsheimiella scutata was located by Barry 
Spracklin. 



LITERATURE CITED 

Eliding, C. 1963. A critical survey of European taxa in Ulvales. 

I. Opera Botanica (Suppl. Bot. Not.). 8: 1-160. Lund. 
Cardinal, A. 1964. Etude sur les ectocarpacees de la Manche. Nov. 

Hedw. 15: 1-86. Ul figs. 

Drouet, F. 1968. Revision of the classification of the Oscilla- 
toriaceae. Monogr. 15. Acad. Nat. Sci. Philadelphia. 



1975] Nahant Algae — Webber 433 



Farlow, W. G. 1882. The marine algae of New England. U.S. 
Comm. Fish and Fisheries. Commissioners Report, 1879. Wash- 
ington: Government Printing Office. Pp. 1-210. 

KOSTER, J. T. 1955. The genus Rhizoclonium in the Netherlands. 

" Puh. Sta. Zool. Napoli. 27: 335-857. 

L.amr, I. M., & M. H. Zimmermann. 1964. Marine vegetation of 
Cape Ann, Essex County, Massachusetts. Rhodora «G: 217-254. 

MATHIESON, A. C, & R. A. FraliCK. 1972. Investigations of New 
England marine algae. V. The algal vegetation of the Hampton- 
Seabrook estuary and the open coast near Hampton, N.H. Rho- 
dora 74: 406-435. 

Stone. R. A., E. J. Hehre, J. M. Conway, & A. C. Mathieson. 

li)70. A preliminary checklist of the marine algae of Campo- 
hello Tsland, New Brunswick, Canada. Rhodora 72: 313-3.°,8. 
Twlor, W. P. 1957. Marine Algae of the Northeastern Coast of 
North America. (Rev. ed.) Univ. Mich. Press. Ann Arbor. 509 
pp., C>0 pis. 

Webber, E. F. 1975. Phycological studies from the Marine Science 
Institute. Nahant, Massachusetts. I. Introduction and prelimi- 
nary tabulation of species. Rhodora 77: 149-158. 

WlLCF, P. T. 1965. Studies in the ^enus Laminaria. "Rot. Gotho- 
bunrenMa. ITT. Proc\ Fifth Mar. Biol. Symp. Gbteborff. 247-256. 



I>i"!FT. OF BIOLOGY 

KKT'KA COLLEGE 

KET T KA PARK, N.Y. 14478 



LACTUCA MURALIS IN MAINE 



James P. Poole 



In a letter dated November 15, 1969, Mrs. Roberta G. 



Mass 



mm 



(L.) Gaertn. on Mt. Desert Island in Salisbury 



Mai 



It was 



collected from a small group of plants that were still in 
good condition when checked again in 1970 and 1971. 
Voucher specimens have been deposited in the Jesup Her- 

Dartmouth College (HNH), in the University 
of New Hampshire Herbarium (NHA), in the Pringle 

at the University of Vermont (VT), and in 



um 



um 



Massachusett 



herst, Mass. (MASS). Collection of this species in Cornish, 
Sullivan County, N.H. was reported in Rhodora 68:774,' 
1966. At that time Professor Albion R. Hodgdon at the 



m 



covery was the first report of the species in New England, 

and Frank C. Seymour's Flora of New England published 

in 1969 lists Cornish as the only New England station. 

Mrs. Poland's discovery represents a really disjunctive 
jump. 



JESFP HERBARIUM 
DARTMOUTH COLLEGE 
HANOVER, N.H. 03755 



434 



LACTUCA MURALIS IN BRITISH COLUMBIA 

John T. Witherspoon and Thomas J. Watson, Jr. 

Lactuca muralis (L.) Gaertn. was recently collected in 



British Columbia (KOA campground, ca. 10 mi W of Hope, 
2 Sept. 1973, Watson 1048). The specimen is deposited at 
MONTU. This is the most inland known record in British 
Columbia as well as the farthest north in the foothills of 
the Cascade Mountains. The locality, latitude 49°20'N 
longitude 121°40'W, is approximately the same longitude 
as a reported locality in the Cascade foothills of Marion 

County, Oregon (Peck, 1961). 

Lactuca muralis is an introduction from northern Europe 
n „A ia oncriTKrlv nHvPTitivP in North America. In eastern 



Q 



New 



York and west to Michigan (Fernald, 1950; Gleason 
and Cronquist, 1963). It was recently reported from New 
Hampshire (Poole, 1966). 



It is apparently more common 



of 



North America and previous to this report it was known 
only from the Oregon locality above, Clallam County and 
San Juan Island, Washington, and from Lake Cowichan. 
Vancouver Island, British Columbia in western North 



Amer 



m 



slowly inland, both eastward and northward in mainland 



Columbi 



LITERATURE CITED 



FERNALD, M. L. 1950. Gray's Manual of Botany. 8th ed. Van 

Nostrand Reinhold Co., New York, lxiv + 1632 pp. 
Gleason, H. A. and A. Cronquist. 1963. Manual of Vascular Plants 

of Northeastern United States and Adjacent Canada. D. Van 
Nostrand Co., Inc., Princeton, li + 810 pp. 

Hitchcock, C. L., A. Cronquist. M. Ownbey and J. W. Thompson. 

1955. Vascular Plants of the Pacific Northwest. Part 5, Com- 
positae, by A. Cronquist. Univ. of Washington Press, Seattle. 

343 pp. 
Poole, J. P. 1966. Lactuca muralis in New England. Rhodora 

68: 138. 



DEFT. OF BOTANY 
UNIVERSITY OF MONTANA 
MISSOULA, MT 59801 



435 



CHIMAPHILA MACULATA IN NEW HAMPSHIRE 



James P. Poole 



In December, 1966, F. S. Page, Emeritus Professor of 
Botany at Dartmouth College, collected two specimens of 
Chimaphila maculata (L.) Pursh on a wooded hill two 
miles south of George Pond and about J/ 2 mile west of 
the dirt road that borders the west shore of the pond. 
The plants bore neither flowers nor fruit, but the leaves 
were sufficiently characteristic for identification. Since in 
Gray's Manual, 8th edition, this species is listed as growing 
in southern New Hampshire only, the collection was ac- 
cessioned in Jesup Herbarium as voucher specimens for 
what appeared to be a range extension for the species. The 
following summer, in August, Professor Page guided Pro- 
fessor Carl Wilson and me to the area where the plants 
had been collected, but careful search over the whole area 
by the three of us failed to turn up another specimen. In 
light of our experience it is interesting to quote from the 
article covering the distribution of this species in Maine 
and New Hampshire by Albion R. Hodgdon and Lesley M. 
Eastman in Rhodora 75:162-165 (1973). "In northern 
New England Chimaphila maculata tends to occur as dis- 
junct colonies of a few individuals or sometimes single 
plants, in this respect resembling some species of orchids. 
Even with careful searching it is usually impossible to find 
any more plants after having found a few shoots at any 
one station. While the stems are of low stature and st«nH 
but a few inches above the forest floor, the leaves 
conspicuous and easy to detect." In the same article a 
number of records extending the range of this species in 
New Hampshire are listed. One collection is listed for 
Tuftonboro, Carroll County, on Sentinel Mountain. 



are 



ap- 



mi 



Grafton County, but I find no record of an earlier collection 
in Grafton County. I am listing this station as in Enfield, 
Grafton County, but it is difficult to locate exactly on the 



436 



1975] Chimaphila maculata — Poole 437 



m 



on either side of the boundary separating Enfield from 
Grantham in Sullivan County. The only other New Hamp- 
shire specimen in Jesup Herbarium was collected on a 
wooded hill in Kensington, Rockingham County, in July 
1896, and bears a printed label reading "Ex Herb. Alvah 
A. Eaton." 



JESEP HERBARIUM 
DARTMOUTH COLLEGE 
HANOVER, N.H. 03755 



BOOK REVIEW: 
WINTER KEYS TO WOODY PLANTS OF MAINE 



i 



When 



taxonomy in the more 
one must, of necessit 



due consideration to those plants that are available during 
the winter season — our woody vascular plants. I enthusi- 
astically welcome this excellent field manual for the identi- 
fication of woody plants in winter condition. The authors 
of this carefully constructed, excellently illustrated work 
have succeeded in providing- a thorough treatment of the 
woody plants of Maine, native, naturalized and adventive, 
with nearly 250 species included. The work goes beyond 
the "clearly" woody species and provides a means for iden- 
tifying such marginally woody species as Arceuthob 
pusillum, Mitchella repens, PotentiUa tridentata, Aralia 
hispida, A. nudicaulis and Decodon verticillatus. The keys 
are well constructed, strictly dichotomous, and usually 



11 HI 



employ a minimum 



Frequently 



mation 



etative features is provided in the leads to further facilitate 
the identification process. Habitat data are also incorpor- 
ated in keys where helpful. The value of the manual is 
greatly enhanced by the superb set of plates of original 
line drawings, made from fresh material. In addition to 
drawings of twigs and of critical leaf scar or bud features, 
the illustrator has provided drawings of leaves and fruits 
of certain species in which they are frequently found per- 
sistent. It is no understatement when the authors claim it 
to be fully illustrated. 

The manual is organized into a "Key to Genera and Aber- 
rant Species" followed by a "Key to Species" with genera 

'Campbell, Christopher S. and Fay Hyland. 1975. Winter Keys 
to Woody Plants of Maine. Illustrated by Mary L. F. Campbell. 
University of Maine Press, Orono, Maine. 52 pp + 63 plates. $3.00. 
Available from: Secretary, Department of Botany, Deering Hall, 
University of Maine, Orono, Maine 04473. 



438 



1975] Winter Keys — Crow 439 



arranged alphabetically. Additional notes are included at 
the end of each set of keys providing information regard- 
ing infraspecific taxa. Nomenclature follows Fernald, 
Gray's Manual of Botany edition 8. A clear, concise and 
complete glossary is also provided. The very thorough 



index 



name, but also lists 



every conceivable combination of common names (e.g., 
Pyrus americana; American Mountain Ash ; Ash, American 
Mountain; Mountain Ash, American). The back inside 
cover provides a map of the counties of Maine and a metric- 
English scale. 

The publication is appropriately dedicated to the mem- 
ory of Professor Merritt Lyndon Fernald, whose long term 
interest in the plants of New England and northeastern 
North America found its stimulus in the plants of his boy- 
hood haunts in the vicinity of Orono, Maine. This fine piece 
of work should be of interest and value to students, ama- 
teur and professional botanists, ecologists, horticulturalists, 
foresters and naturalists of not only Maine, but of her 
adjacent states and provinces. At $3.00, this is a welcome 
bargain. 



Garrett E. Crow 

DEPARTMENT OF BOTANY & PLANT PATHOLOGY 
UNIVERSITY OF NEW HAMPSHIRE 
DURHAM, NEW HAMPSHIRE 03824 



Volume 77, No. 811, including pages 337-U39, was issued Dec. 15, 1975 



Cover illustration 

Rhododendron canadense (L.) Torr 

from 
Curtis' Botanical Magazine t.474 



RHODORA September, 1975 Vol- 77, No. 811 



CONTENTS 



A Revision of the North American Genus Amorpha (Legummosae- 
Psoraleae). 

Robert L. Wilbur 337 



410 



Papaver in North America North of Mexico. 
Robert W. Kiger 

The North American Species Boerhavia s pic at a (Nyctaginaceae), 
in Northwestern Argentina. 
Philip D. Cantino , 423 

Geographical Distribution of the Cedar Glade Endemic Viola 
egglestonii. 

Jerry M. Baskin and Carol C. Baskin 427 

Recent Additions to the Marine Algal Flora of Nahant, Massa- 
chusetts. 

E. E. Webber 430 



Lactuca muralis in Maine. 
James P. Poole 

Lactuca muralis in British Columbia. 



434 



John T. Witherspoon and Thomas J. Watson, Jr 435 



436 



Chimaphila maculata in New Hampshire. 
James P. Poole 

Book Review: Winter Keys to Woody Plants of Maine. 
Gcurrett E. Crow „...„ 438 




JOURNAL OP THE 



NEW ENGLAND BOTANICAL CLUB 




Vol. 77 



December, 1975 



No. 812 



QUfe ^tfa ^ttglanh ^otouotl Club, $nt. 



Botanical Museum, Oxford Street, Cambridge, Mass. 02138 



Conducted and published for the Club, by 
ALFRED LINN BOGLE. Editor-in-Chief 

HOLLA MILTON TRYON 
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Associate Editors 



RHODORA. — A quarterly journal of botany, devoted primarily to 
the flora of North America and floristically related areas. Price 
$20.00 per year, net, postpaid, in funds payable at par in the United 
States currency at Boston. Some back volumes, and single copies 
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plants of North America, will be considered by the editorial com- 
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and cytotaxonomy in their broader implications are equally accept- 
able. All manuscrints shnnM " 



be 



be 



double-spaced throughout. Please conform to the style of recent 

issues of the journal. Illustrations can be used only if the cost of 

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Forms may be closed five weeks in advance of publication. Ex- 
tracted renrints. if nrdprod in advanoa «.;n u * » . « , , 



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Pathology 



03824 



Subscriptions and orders for back issues (making all remittances 
payable to RHODORA) should be sent to RHODORA. Botanical 
Museum, Oxford Street, Cambridge, Mass. 02138. In order to receive 
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Second Class Postage Paid at Boston, Mass. 



MANUFACTURED BY 
THE LEXINGTON PRESS, INC. 
LEXINGTON, MASSACHUSETTS 



Cover illustration 

Rhododendron canademae (L.) Torr, 

from 
Curtis' Botanical Magazine t.474 



IRbofcora 



JOURNAL OF THE 



NEW ENGLAND BOTANICAL CLUB 



Vol. 77 December, 1975 No. SI 2 



HYBRIDS IN AMERICAN CYATHEACEAE 



David S. Conant 



Several small tree ferns collected in a ravine near Cerro 
de Punta, Puerto Rico, are proposed as a hybrid between 
Alsophila dryopteroides (Maxon) Tryon and Nephelea 
portoricensis (Kuhn) Tryon, both of which were growing 
abundantly at the site. This is the first unequivocal report 
of a hybrid in the family. 

In addition, six other hybrids are proposed, one on the 
basis of substantial evidence, and the others on the basis 
of inference. Four previous reports suggest that hybridiza- 
tion may occur in the Cyatheaceae, but definitive analyses 
of the parental and hybrid characters were not presented. 
Holttum and Sen (1961) and Holttum (1963, 1974) report 
that Cyathea alternans (Wall ex Hook.) 



M 



Sumatra and Borneo might be a series of hybrids between 
Cyathea moluccana R. Br. and C. squamulata (Bl.) Copel. 

itecture. Gastony 



dusium 



(1973) treats Nephelea concinna (Kuhn) Tryon as a spe- 
cies but notes the possibility that it is a hybrid between 
N. pubescens (Kuhn) Tryon and N. Tussacii (Desv.) 
Tryon on the basis of intermediate characters of the lamina 
architecture and partially contracted fertile pinnules. 



441 



442 Rhodora [Vol. 77 



s 



DEFINITE HYBRIDS 

1. Alsophila dryopteroides (Maxon) Tryon X Nephelea 
portoricensis (Kulin) Tryon 

■ 

Plants of this hybrid (Fig. 1) resembled large plants of 
Alsophila dryopteroides at a distance, but on closer inspec- 
tion they were seen to have the black squaminate spines of 
Nephelea on the petiole. Species of Cyatheaceae growing 
nearby were Lophosoria qnadripimmta (Gmel.) C. Chr., 
Cyathea arborea (L.) Sm., C. furfuracea Baker, Trichip- 
tcris armata (Sw.) Tryon, T. borinquena (Maxon) Tryon, 
Alsophila, hryophila Tryon, A. dryopteroides (Maxon) 
Tryon, and Nephelea portoricensis (Kuhn) Tryon. 

The black squaminate spines on the petiole and rachi 
(Fig. 10) are a generic character of Nephelea (Fig. 12). 
They firmly establish one parent as N. portoricensis since 
it is the only member of the genus in Puerto Rico. The 
setate scales on the lamina of the hybrid restrict the second 
parent to a species of Alsophila of which there are three 
in Puerto Rico; A. Brooksii (Maxon) Tryon, A. hryophila, 
and A. dryopteroides. Alsophila Brooksii is not known 
from the central mountains where the hybrids grew, it 
lacks the dark lanceolate type of scale present in the hybrid, 
and its long petiole is inconsistent with the requirements 
for the second parental species. Similarly, A. hryophila 
may be eliminated as a possible parent because it also lacks 
the dark lanceolate type of scale, and a parent with a short 
rather than a tall trunk is required. Alsophila dryopter- 
oides remains as the second possible parent and all of the 
characters of the hybrid are consistent with this choice. 

A detailed comparison of the hybrid and parental spe- 
cies is presented in Table 1. Some of the 

acters of the hybrid are in the leaves, stems, and lamina 
scales. 

Characters which are intermediate are lamina shape and 
dissection, stem, petiole and rachis spininess, number of 
leaf scars per unit of stem length, and leaf scar diameter. 
The lance-ovate lamina of the hybrid is intermediate be- 



impor 



1975] 



Cyatheaceae — Conant 



443 





















' 




Figs. 1-2. Alsophila dryopteroides X Nephelea portoricetnsis : 

1, plant, stem ca. 75 cm. tall, leaves ca. 1.0 m. long, Conant G79; 

2, spore, X 1500, Conant 687. Figs. 3-4. Cnemidarui horrida X 
Cyathea arborea: 3, plant, stem ca. 2.0 m. tall, leaves ca. 2.5 m. 
long, Conant 626; 4, bicolorous scales on crozier, X 1/3, Conant 54(5. 
(all specimens GH). 



444 



Rhodora 



[Vol. 77 






. . 






* 



5 




Nephelea 



D 













Aisophiia 



hybrid 







%w 













5, leaf tracings, A, N. por- 



Figs. 5-6. Leaf tracings and pinnae of Nephelea portoricensis, 
Hybrid, and Alsophila dryopteroides: 

toricensis, B, from Conant 679 and C, from Conant 680, Hybrid, 
D, from Conant 681, E, from Conant 690 and P, from Conant 682, 
A. dryopteroides, 



X 



1/30; 



portor 



c&ms, B, Conant 1980, Hybrid, C, Conant 1852, A. dryopteroides, 
X 1/5. (all specimens Gh). 



1975] Cyatheaceae — Conant 445 



mm 



tween the oblanceolate lamina of A. dryopteroides and the 
ovate lamina of N. porta rice n sis (Fig. 5). The lamina 
dissection of the hybrid is bipinnate-lobed which is inter- 
mediate between the pinnate-pi nnatifid condition of A. 
dryopteroides and the bipinnate-pinnatifid condition of N. 
portoricensis (Fig. 6). The stem, petiole and rachis of the 
hybrid have a few spines reaching 2.5 mm. long (Figs. 9 
and 10), a condition which is intermediate between the 
spineless A. dryopteroides (Figs. 7 and 8) and the numer- 

long in N. portoricensis (Figs. 

11 and 12). The hybrid has from 18-25 leaf scars 1.5 cm. 

of stem (Fig. 9). Alsophiki dry- 
opteroides has about 43 leaf scars 1.0 cm. in diameter, per 
10 cm. of stem (Fig. 7) while N. portoricensis has about 
13 leaf scars 2.0 cm. in diameter, per 10 cm. of stem 
(Fig. 11). 

One character of the hybrid is similar to that of Al- 
sophila dryopteroides. The dark lanceolate laminar scales 
with a single apical seta found in the hybrid (Fig. 14) 
closely resemble the most prominent laminar scale of A. 
dryopteroides (Fig. 13). 



am 



Neph 



portoricensis. 



lami 



N 



and the marginally setate type of scale also occurs in the 
hybrid (Fig. 16) and in N. portoricensis (Fig. 18). 

The spores of the hybrid plants are well developed 
(Fig 2). A study of their viability has been planned but 
until it is completed it is not possible to suggest the number 
of F, hybrids probably involved in the five plants seen. 

Trunk erect, occasionally short spiny, to 2 m. high, leaf 
scars 1.5 cm. in diameter, 18-25 per 10 cm. of stem length. 
Leaves 1.0-1.5 m. long, petiole to 10 cm., armed with short 
black squaminate spines; lamina lance-ovate, gradually nar- 
rowed at base (basal pinnae to \\ as long as the median), 
to 60 cm. broad at the middle, acuminate, bipinnate-lobed 
to occasionally bipinnate-pinnatifid, leaf tissue glabrous, 



446 



Rhodora 



[Vol. 77 



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Figs 



r'cetixia. 



7-18. Ahophila dryopteroides, Hybrid, and Neplielea porto- 
Figs. 7-8. A. dryopteroides: 7, stem, X 2 3, Conant (582, 
8. petiole, X 2, Comitwi 690. Figs. 9-10. Hybrid: 9, stem, X 2 3, 
CottrtW* 680, 10, rachis, X 2, Cowu// 680. Figs. 11-12. N. portori- 
eensift: 11, stem, X 2 3, 7Y//<m & 7Y.i/<m 6950, 12, petiole, X 2/3, 



Conant 412. Fi 






1 ° 

I O. 



A. dryopteroides, lanceolate type scale, X 100, 
Conant 598. Figs. 14-16. Hybrid: 14, lanceolate type scale, X 100, 
Cowanf 687, 15, acaroid scale, X 100, Conant 687, 16, marginally 
set.ate scale, X 100, Conant 687. Figs. 17-18. N. portoricensis : 17, 
acaroid scale, X 100, Conant 188, 18, marginally setate scale, X 100, 
Conant 188. (all specimens GH). 



448 Rhodora [Vol. 77 



herbaceous, glaucescent beneath, dark green above; rachis 
dull castaneous, occasionally with short spines at the base, 
glabrescent to squamulose beneath, invested with stiff, 
brown, antrorse trichomes and a few long filiform scales 
above; pinnae to 25, spreading, sessile, the largest 30 cm. 
long, 7 cm. broad near base, becoming pinnatifid towards 
the apex; pinna-rachis beneath invested with marginally 
setate scales, acaroid scales, and dark bodied, light mar- 
gined, lanceolate scales with a single, dark, stout apical 
seta, and with intermediate scales ranging from oblong to 
lanceolate, uniformly brown to dark bodied, and many 
setate to singly and stoutly setate, these latter types be- 
coming sub-bullate towards the apex; undersurface of pin- 
na-rachis becoming sparsely pubescent apically; pinnules 
18-24 pairs, 19-34 mm. long, 7-10 mm. broad, deeply lobed 
to occasionally pinnatifid at the base; costa beneath often 
with acaroid scales at the base, and beyond with sub- 
bullate scales with few to many setae, these progressing 
to bullate .scales at the pinnule apex, occasionally with a 
few stiff trichomes apically; costules with marginally setate 
sub-bullate scales at the base, these grading into squamules 
apically; veins 4-5 pairs, sori 2-3 on each lobe, closer to the 
costule than to the margin ; indusia deeply cyathiform, tan, 
membranous, glabrous; spores well developed, light tan. 
trilete. 



SPECIMENS EXAMINED: 

Puerto Rico: Five plants from the vicinity of Cerro de Punta, 
40 miles w.s.w. of San Juan, ravine on north side of road, Rt. 143, 
km. 18.5 e. of jet. with Rt. 10, Conant ()78 (gh, ny, us), Co'nant 
679 (GH, NY, us), Conant 680 (gh, ny, us), Conant 687 (F, gh, ij, 
NY, rppr, us, USD, Herb. El Verde Field Station, Puerto Rico) ; 
ravine on north side of road, Rt. 143, km. 22.2 e. of jet. with Rt. 
10, Conant 599 (gh). 



2. Cnemidaria horrida (L.) Presl X Cyathea arborea (L.) 
Sm. Hemitelia Wilsonii Hook, in Hook. & Baker, Syn. 
Fil. 30. 1865. type: Jamaica, Wilson 731 (k). (See 
Maxon, Contrib. U.S. Nat. Herb. 17: t. 18. 1914.) 



1975] Cyatheaceae — Conant 449 



The status of Hemitelia Wilsonii (Fig. 3) as a species is 
untenable because well developed spores have not been 
found in its sporangia. Other factors indicating a hybrid 
status are the highly variable leaf architecture and the 
variable venation. 

I have seen Hemitelia Wilsonii growing at two sites in 
the Luquillo Forest, 20 miles e.s.e. of San Juan, Puerto 
Rico (Rt. 186, km. 19.0 s. of jet. with Rt. 3, Municipio de 
Rio Grande, and Sabana Rd., km. 3.0 e. of jet. with Rt. 191, 
Municipio de Rio Grande). In both places Cnemidaria 
horrida and Cyathea arborea were growing nearby. 

The hybrid is intermediate between the parental species 
in its venation, indusia, and leaf architecture. The venation 
of the hybrid is usually free but occasionally anastomoses 
or forms loops or dead-ends or both (Fig. 22). This con- 
dition is intermediate between the free venation of Cyathea 
arborea (Fig. 24) and the anastomosing venation of Cne- 
midaria horrida (Fig. 20). The indusia of the hybrid range 
from hemiteloid to sub-cyathiform (Fig. 21), a condition 
intermediate between the hemiteloid indusia of Cn. horrida 
(Fig. 19) and the cyathiform indusia of C. arborea (Fig. 
23). The leaf architecture of the hybrid ranges from 



nnate 



m 



ate between the pinnate-pinnatifid condition of Cn. horrida 
and the bipinnate-pinnatifid to tripinnate architecture of 
C. arborea. 

Characters of the hybrid which are closest to Cnemidaria 
horrida are the shape of the pinna and leaf apex and the 
bicolorous scales on the croziers and petioles (Fig. 4). 
One character of the hybrid which is found in Cyathea 



small 



on the costa undersurface. 



combinat 
yf Heniitt 



Wilsonii 



support 



H. Wilsonii 



horrida and Cyathea arborea. 



450 



Rhodora 



[Vol. 77 



20} 




Figs. 19-24. Cnemidaria horrida. Hybrid, and Cydthea arborea. 
Figs. 19-20. Cn. horrida: 19, indusia, X 20, Gaxtony et al. 655, 
20, venation, X 7*4, Scamman 8117. Figs. 21-22. Hybrid: 21, 
indusia, upper arrow, subcyathiform indusium, lower arrow, hemi- 
teloid indusium, X 20, Gastouy et al. (r>4, 22, venation, upper arrow, 
anastomosing veins, middle arrow, space between free veins (note 
loop in lower vein), lower arrow, dead-end in vein, X 7*^, Conant 
540. Figs. 23-24. C. arborea: 23, indusia, X 20, Howard & Nevling 
15776, 24, venation, X 7%, Gastony 10. (all specimens Gil). 



1975] Cyatheaceae — Conant 




SPKCIMENS EXAMINED: 



Puerto Rico: Conant 546, 626, 627 (GH), Hess 371 (NY), Hioram 
182, 804 (NY), Kepler, Sabana Rd., Kl Yunque Rd., 5 Mar., 1970, 
Herb, El Verde Field Station, Municipio de Rio Grande. Dominican 
Republic: Abbott 2660 (GH, NY, us), Ekman 14752 (ny), L5014 
(GH, ny), Gastony, Jones & Norris 654 (<;n). Haiti: Ekman H4835 
(NY, s), H4846 (s, US). Jamaica: Jewman (ny) , Proctor 18419 (a). 



PUTATIVE HYBRIDS 

The establishment of the Alsophila-Nephelea intergeneric 
hybrid with a bipinnate-lobed leaf architecture helps to 
interpret other rare species and unusual collections with a 
similar architecture within the Alsophila-Nephelea evolu- 
tionary line. These are discussed below in order of the 
certainty of their parental species. All of them combine 
characters of species with very different leaf architecture 
and (or) lamina indument. All are very rare, known 
mostly from one collection, and do not fit within the vari- 
ation of the species proposed as parents. Nephelca con- 



vex 



N 



(1973) but a detailed 



analysis of it has not been made. 



3. Alsophiln Brooksii (Maxon) Tryon X Nephelea portori- 
censis (Kuhn) Tryon 

A collection from Indiera Fria, near Maricao, Puerto 
Rico (Britton, Cowell, & Brown 4520 (NY) ) resembles the 
AUophila dryopteroides - Nephelca portoricensis hybrid 
quite closely in leaf dissection and in width to length pro- 
portions of the pinnae and pinnules. It differs in that it 
lacks the dark bodied, light margined, lanceolate scales 
with a single, dark, stout apical seta of A. dryopteroides. 
In addition the lamina base is abruptly narrowed, and the 
petiole is up to 25 cm. long. The complexity of the lamina, 
the abundance of acaroid scales, and the pubescence of the 
veins and costules of this collection are all characters 
attributable to N. portoricensis as pointed out by Gastony 
(1973). Thj unusually long and narrow pinnae .suggest 

fi«Qf -Hiia mnv V»p a hvhrid involviuo- N. nortoriceHsis and 



452 Rhodora [Vol. 77 



an Alsophila species with a less dissected lamina and long 
narrow pinnae. Alsophila Brooksii is well known from the 
Maricao Forest about 10 km. to the west of Indiera Fria. 
It is the only species of Alsophila occurring in the western 
end of the island, and it has a pinnate-pinnatifid lamina 
abruptly reduced at the base. It has long narrow pinnae 
and its petiole reaches 78 cm. in length. Its characters, 
combined with those of N. portoricensis, would result in a 
plant very similar to the Btitton et al. 4520. 



4. Nephelea balanocarpa (D. C. Eaton) Try on X Nephelea 
woodwardioides (Kaulf.) Gastony 

Cuban material with bipinnate-lobed lamina architecture 
such as Leon, Clement, & Roca 10533 (ny) and Hioram & 
Clement 6377 (us) seems to represent hybrids between 
pinnate-pinnatifid and bipinnate-pinnatifid species. One 
parent is probably the bipinnate-pinnatifid Nephelea wood- 
wardioides. It is not clear whether var. woodwardioides or 
var. cubensis (Maxon) Gastony is involved. Nephelea 
balanocarpa is probably the second, pinnate-pinnatifid 
parent. It is far more common than A. Brooksii, also pin- 
nate-pinnatifid, which is known in Cuba only from the type 
collection. The abundance of tiny antrorse squamules cov- 
ering the veins in the hybrid is in agreement with the in- 
dument of N. balanocarpa. 



5. Alsophila hotteana (C.Chr. & Ekman) Tryon X Nephe- 
lea sp. Cyathea confinis C. Chr. Kungl. Svensk. Vetens.- 
akad. Hand. ser. 3, 16:13. 1937. type: Haiti, Massif de 
la Hotte, Jeremie, Ekman 10382 (holotype, s; isotype, 

us!). 

Cyathea confinis, a Haitian species known only from the 
type collection at Massif de la Hotte, has a bipinnate-lobed 
lamina, suggesting that it is probably of hybrid origin. 
It is described as having an aculeate trunk to 2 m. high, a 
short petiole armed with short black spines, and a grad- 
ually reduced lamina base. A relationship to the genus 



1975] Cyatheaceae — Conant 453 



Nephelea is evident in the aculeate trunk and the black 
petiole spines. Three species of the #enus occur in Haiti, 
N. crassa (Maxon) Try on, N. fulgens (C.Chr.) Gastony, 
and N. woodwardioides (Kaulf.) Gastony var. Hieronymi 
(Brause) Gastony. All are tall and have a bipinnate- 
pinnatifid lamina making it difficult or impossible to deter- 
mine from herbarium specimens which of the species is 
involved in the cross. 

Alsophila hotteana is probably the pinnate-pinnatifid 
parent. Its short petiole, gradually narrowed lamina base, 
and tall stem are consistent with the characters required 
of the second parent, and it is the only Alsophila known to 
occur in the Massif de la Hotte. Other species of Alsophila 
in Haiti are restricted to the Massif du Nord. 



Putative Hybrids Involving Undetermined Species 

of Alsophila and Nephelea 

6. Cyathea irregularis Brause Urban Symb. Ant. 7. 155. 
1911. TYPE: Dominican Republic, Santo Domingo, Con- 
stanza, Turckheim 3212 (holotype, B; isotype, ny!). 

Cyathea irregularis, known only from the type collection, 
is another species with a bipinnate-lobed lamina. It is evi- 
dently a hybrid of Alsophila and Nephelea since all species 
of Alsophila in the Dominican Republic are pinnate-pin- 
natifid and all species of Nephelea are bipinnate-pinnatifid. 
The single specimen of a midportion of the lamina, how- 
ever, provides too few characters to choose between the 
four Alsophilas and three Nepheleas known to grow in the 
country. 

Another specimen from the Dominican Republic, with a 
similar leaf morphology, Abbott 2031 (us, pro parte) is 
probably an Alsophila X Nephelea hybrid but may not 
have the same parents as Cyathea irregularis. 



7. Cyathea jamaicensis Jenm. Jour. Bot. 20: 323. 1882. 
TYPE: Jamaica, Mansfield, near Bath, Wilson 686 (holo- 
type, bm; isotype, gh!). 



454 Rhodora [Vol. 77 



The bipinnate-lobed Cyathea jamaicensis is known only 
from the type collection. It is similar to Nephelea con- 
cinna, but may be separated from it by its pinna rachis 
indument and by its indusium. Cyathea jamaicensis has 
the undersurfaces of its pinna-rachises nearly without 
scales and trichomes and it has a meniscoid indusium. 
Nephelea concinna has its pinna-rachis undersurfaces quite 
well invested with scales and trichomes and it has a cyathi- 
form to urceolate indusium. Also, C. jamaicensis is de- 
scribed as having" an unarmed stem whereas N. concinna 

has a spiny stem. 

It seems probable that Cyathea jamaicensis is a hybrid 
because of its lamina architecture and its rarity. The lack 
of spines on its trunk implies that it could not be an intra- 
generic cross involving two species of Nephelea. The de- 
gree of lamina dissection of C. jamaicensis suggests that a 
bipinnate-pinnatifid species of Nephelea has crossed with 
a pinnate-pinnatifid species of Alsophila. 



CONCLUSIONS 



Two certain and five putative hybrids have been pro- 



posed. 



}f species with very differ- 
dument, and (or) indusia. 



The implication is that other species with similar leaf 
architecture, lamina indument, and (or) indusia should 
also cross. These hybrids would be very difficult or im- 
possible to detect. Some polymorphic species in current 
taxonomie treatments may require reassessment because 



may 



Careful field 



observations of species that are growing together, when- 



ever tree fern collections are made, will help to solve this 
problem. 

The presence of well developed spores in the Alsophila 
dryopteroides X Nephelea portoricensis hybrid suggests 
that these genera are closely related. The apparent absence 
of well developed spores in the Cnemidaiia horrida X 
Cyathea arborea hybrid implies a more distant relation of 
the parents. 



1975] Cvatheaceae — Conant 455 



ACKNOWLEDGEMENTS 



The writer wishes to express sincere thanks to the Puerto 
Rico Nuclear Center, U.S. Atomic Energy Commission, 
Rio Piedras, Puerto Rico, and particularly to Dr. Richard 
G. Clements, Director, Division of Terrestrial Ecology, for 
making Student Research Grants available during 1972 
and 1973 when collections and field studies were made. 
Dr. Rolla M. Tryon has contributed to the organization and 
presentation of ideas and conclusions embodied in this 
paper. Thanks are due to Dr. Alice F. Tryon and Paulo G. 
Windisch for help in preparing the plates and for percep- 
tive discussion. I am indebted to Bruce Tiffney and to 
Dr. Elso Barghoorn for assistance in the use of the photo- 
graphic equipment of the Paleobotanical Laboratory at 
Harvard. Many thanks go to Jose and Elvira Colon, and 
to Michael Burke for assistance in making collections and 
in pressing specimens. 



LITERATURE CITED 



Gastony, G. J. 1973. A revision of the fern genus Nephclea. 

Contrib. Gray Herb. 203: 81-148. 
Holttum, R. E. 1963. Cvatheaceae. Flora Malesiana II, 1(2): 



65-176. 



— . 1974. The tree ferns of the genus Cyathea in Borneo. 
Garden's Bull. 27: 167-182. 
, & U. Sen. 1961 Morphology and classification of the 



tree ferns. Phytomoi phology 11: 406-420. 



BIOLOGICAL LABORATORIES 
HARVARD UNIVERSITY 
CAMBRIDGE, MASS. 



ZOSTERA MARINA L., ITS GROWTH 

AND DISTRIBUTION IN THE 
GREAT BAY ESTUARY, NEW HAMPSHIRE 



Stanley A. Riggs, Jr. and Richard A. Fralick- 



Protected coastal and estuarine waters of New England 
often contain dense meadows of the marine Angiosperm 
Zostera marina L. Little was known of the ecology of 
Zostera, or eel grass as it is commonly called, until a wast- 
ing disease attributed to the parasitic marine fungus Laby- 
rinthula almost exterminated the Zostera population in 
New England waters (Jepps, 1931; Renn, 1934, 1935, 
1936a, 1936b; Young, 1937, 1943). 

The present paper compares the seasonal growth (bio- 
mass), reproductive phenology, and local distribution of 
Zostera with seasonal variations in temperature and salin- 
ity. The results of vertical and horizontal transplants are 
discussed as well as the possibility of ecotypical adaptation. 



METHODS AND MATERIALS 



Bimonthly observations and measurements of Zostera 
marina populations were made at eight stations within the 
Great Bay estuary system during 1972 (Table 1). Meas- 
urements of surface water temperature and salinity were 
made at mean low water (MLW). Biomass was also re- 
corded at MLW since the majority of Zostera plants oc- 
curred in this zone. 



'Jackson Estuarine Laboratory Scientific Contribution Number 27. 

2 Present address: Department of Natural Science, Plymouth State 
College of the University of New Hampshire, Plymouth, N.H. 03264. 



456 



1975] 



Zostera — Riggs & Fralick 



457 



Table 1, Characteristics of eight stations within the 

Great Bay Esturary. 



Stations/ 

nautical 

miles inland 



1. Jaffrey Point 
(0.0) 



Temperature 

°C 



avg. 

max. 

min. 



10.2 
23.0 

1.0 



Salinity 



Substrate 



V 



00 



avg. 
max. 



30.3 rocky outcrops 
32.5 with some sand 



min. 26.0 



2. Pierce Island 
(2.0) 



avg. 
max 
min. 



10.5 

21.5 

0.0 



avg. 27.0 broken rocks, mud 
max. 32.0 and silt 



min. 



19.0 



3. Newington Town avg 



Landing (5.5) 



max 
min. 



11.7 

24.0 

0.4 



mm. 



24.5 small rocks, mud 



avg. 

max. 31.0 and silt 



16.0 



4. Dover Point 
(7.0) 



avg. 
max. 

min. 



12.5 

25.0 

3.0 



avg. 23.7 large rock outcrop- 
max. 32.0 pings, mud and silt 
min. 15.0 



5. Cedar Point 
(8.3) 



avg. 



13.6 



max. 27.0 
min. 0.0 



avg. 22.1 mud and silt 



max 
min. 



32.0 
6.0 



6. Adams Point 
(10.7) 



avg. 13.8 
max. 28.0 

min. 0.5 



avg 



21.0 rock outcroppings. 



max. 31.5 shale, cobble, 
min. 8.0 mud and silt 



7. Chapmans 
Landing (15.2) 



avg. 
max 



min. 



14.2 

26.5 

0.0 



avg. 5.7 mostly mud, 
max. 22.0 some small 



mm. 



0.0 rocks and boulders 



8. Exeter 
(19.1) 



avg. 14.4 

28.1 



max, 
min. 



0.0 



avg. 
max. 

min. 



3.4 mud and silt 
20.0 
0.0 



458 Rhodora [Vol. 77 



Growth studies were initiated during March, 1972. 
Tagged plants (approximately 100) were established at 
Stations 2 through 6, and the blade length was measured 
according to the method of McRoy (1970). Horizontal and 
vertical transplants of Zostera were also initiated in March, 
and the plants were subsequently observed and measured 
for a period of 10 months. Each of the transplants was 
duplicated, one using the substrate of the original station 
and the other using the substrate of the new station. 

McRoy (1970) showed that a rapid estimate of the stand- 
ing crop of Zostera is possible because of a relationship 
between blade length and dry weight. By incorporating 
both the width and the length of the blades we derived and 
untilized the following equation for conversion to dry 
weight (biomass) without harvesting or damaging the 
plants. 



Formula Example : 

Y = (0.308 X 10" 4 ) (x) - 48.2 X 10 4 

Y = dry wt. (grams) 
x = L X W (mm) 

blade width = 5 mm 
blade length = 100 mm 

L X W = 500 
Y = (0.308 X 10 4 ) (500) - 48.2 X 10 4 
predicted - 10.6 X 10 3 g dry wt. 
actual - 10.8 X 10 3 g dry wt. 



A linear regression was performed between the predicted 
dry weight and the actual dry weight, and a correlation co- 
efficient of 0.99 was obtained. 

Temperature was recorded with a laboratory grade, sub- 
mersible, mercury thermometer. Salinity was determined 
with a S3t of hydrometers (G.M. Manufacturing Co., N.Y.) 
and the readings were corrected to 15°C. 



1975] 



Zostera — Riggs & Fralick 



459 



Hydrographic factors: Figure 1 shows the seasonal vari- 
ation in temperature at two representative stations for 12 
months. Stations 1 and 2 exhibited similar temperature 
values with maximum temperatures of 23.0° and 21.5° C 
occurring in July and minimum temperatures 1° and 0° C 
occurring in January, February, and December. Stations 

4 exhibited maximum temperature values of 25.0° 
to 26.0° C in July with minimum values of 0° C during the 
winter months. Stations 6, 7, and 8 showed the greatest 
ranges in temperature with maximum values of 27.5° C 
in July and minimum values of 0° C in January, February, 
and December. 



8 and 



u 



Q£ 



< 



s 



i-in itial Growth 



FIG. 1 



STATION 2 



STATION 




MONTHS (1972) 



460 



Rhodora 



[Vol. 77 



Figure 2 shows the seasonal variation of salinity at four 
representative stations for 12 months- The values are con- 
sistent with the proximity of each station to the open coast. 
Thus, Station 1 had a seasonal salinity range of 26.0 to 
32.5 o/oo. ; Station 2 showed a range of 19.0 to 32.0 o/oo. ; 
Station 6 had a range of 8.00 to 21.0 o/oo., while Station 8 
had the lowest salinities within a range of 0.0 to 20.0 o/oo. 



M 





< 

CO 



i-lnitial Growth 
f-Flowering 

b-Biomass Max 



STATION 2 
STATION 3 



F 




M 



A 



M 



J 



J 



A 



S 



O 



N 



D 



MONTHS 



J 



Seasonal growth: Figure 3 indicates that active growth 
of Zostera was initiated during March and April at Sta- 
tions 2, 3, 5, and 6. The plants at Stations 2 and 3 exhibited 
a conspicuous elongation of the blades (1 to 3 cm) in 
March. The plants at Stations 5 and 6 did not initiate 
growth until the end of April. Growth at Stations 2, 3, 5, 
and 6 declined rapidly during November and December 
and no growth was recorded in January and February. No 



1975] 



Zostera — Riggs & Fralick 



461 



Q ° 




< 



I • 



FIG. 2b 



5f ;a\ c , / 

U\ '■■■ V ••• 

6f\ / 



6 b ... 



\ V 



1 / «5i fl i 

J / if: * \6i v 







\ 



I 



i 



w 



\ 



; 



I 

/ ! 
/ ••• 






i-lnitial Growth 
f -Flowering 
b-Biomass Max 



STATION 5 

STATION 6' 



MONTHS (1972) 



growth was observed at Stations 1, 7, and 8 throughout the 
season. The growth at Station 4 is not illustrated because 
the colony was considered subtidal, while all other stations 
were intertidal. 



Standing crop: Figure 3 illustrates the seasonal varia- 
tion in standing crop of Zostera at several stations. Mini- 
mum values were apparent from December through March. 
Maximum standing crop values were recorded in August 
at Station 5. The plants at Station 6 reached their 



maxi 



mum 



biomass in late September. Two peaks were evident 
at Stations 2 and 3 ; the first occurred in mid-June and the 



lomass 



second in early August. The initial peak in 1 

associated with flowering, while a decrease in early July 

was attributed to the abscission of floral parts. In some 



462 



Rhodora 



[Vol. 77 



FIG. 3 



STATION 

STATION 3* 

STATION 



STATION 6* 



o 



X 

E 
o 

O 

o 

E 
o 




MONTHS (1972) 

cases vegetative turions without reproductive parts re- 
mained intact and continued to grow. 

A general decline in standing crop was apparent at 
nearly all stations by late September, and although some 
growth continued until December, no significant increase 

was recorded at any station after October. The 



omass 



process of floral abscission was not apparent at Station 6 
until early October. 



Horizontal trans 



Transplants were also initiated 



between various stations during the spring. In all cases the 



maximum 



same time as the original populations from which they 
were derived. For example, Zostcra transplanted from 



maximum 



mi 



reached a 



September (Fig. 4). Zostera at Station 4 



maximum 



value in July. 



1975] 



Zostera — Riggs & Fralick 



463 



30 



FIG. 4 



25 



2 20 

X 

E 



o 

o 

E 
o 

CO 



F 



M 



STATION 2 
STATION 4 



STATION 6 



intertidal to subtidal (STA. 6) •- 




A 



M 



J J 

MONTHS 



A 



S 



O 



N 



D 



J 



Transplants of Zostera plants 



cm loner) from 



Station 2 to 8 died within a week. Differential responses 
to transplanting were observed when smaller plants (2.0 
to 8.0 cm long) survived a transplant between the same 
stations for more than 30 days. 



DISCUSSION 



Within the Great Bay Estuary 



tern, Zostera first 
maximum biomass 



at the stations nearest the coast. 



further 



maxi 



mum biomass some three months later tnan stations ciose 

to the coast. 

The initial growth of Zostera, in contrast to Setchell's 

1929 findings, is neither restricted to a specific 5 degree 



464 



Rhodora [Vol. 77 



temperature isotherm, nor to temperatures greater than 
10° C. Although Zostera has been observed living beneath 
winter sea ice (McRoy, 1969), to our knowledge its growth 
has not previously been measured at temperatures below 



10° C. 



maximum biomass 



to be related to any specific temperature range. Thus, 
plants at Stations 2 and 3 reached their maximum biomass 
at 18° C, while it was attained at 25° C at Station 5 and 

16° C at Station 6. 

The flowering period is not restricted to a specific tem- 
perature isotherm (15° to 20° C) as suggested by Setchell, 
1929. Zostera flowered in the spring at Stations 2 and 3 



temp 



° C. At Station 6 



ace 



in temperature up to 28.0° C. 



biomass 



high salinity at each site. Thus, Zostera plants at Stations 
2 and 3 reached their first maximum biomass at salinities 
of 26.0 o/oo and 24.0 o/oo respectively, while maximum bio- 
mass at Station 5 was reached at a salinity of 25.0 o/oo — 
40 days later than at Station 2. Zostera plants at Station 6 
reached maximum biomass at a salinity of 27.0 o/oo, in 
September, nearly 100 days later than at Stations 2 and 3. 
The time of flowering was also correlated with salinity. 
Thus, Zostera plants at Stations 2 and 3 initiated flowering 
when the salinities were 26.0 and 24.0 o/oo. The plants at 
Stations 5 and 6 also initiated flowers at the same salini- 
ties, but approximately three weeks later than at Stations 

2 and 3. 

It appears that Zostera marina maintains its original 

growth patterns even after being transplanted to new loca- 
tions. Thus, transplants from Stations 2 and 4 to 6 fol- 
lowed a pattern of growth and development similar to that 
of the plants in their original colony. In no case did trans- 
planted Zostera follow a sequence of growth and pheno- 
logical development parallel to patterns in plants originat- 
ing in the new location. 



1975] Zostera — Riggs & Fralick 465 



The horizontal transfer of Zostera colonies within the 
estuary had no noticeable effects on their development. 
Vertical transplants (intertidal to subtidal) frequently 
showed a decrease in their rate of development. Thus, 
when young seedlings (1 to 3.0 cm long) were transplanted 
vertically their development was conspicuously slowed 
(Fig. 4). In contrast, transplants from the subtidal to the 
intertidal zone were consistently successful. Transplants 
to brackish waters with salinities of 4.0 to 5.0 o/oo did 
poorly. The greatest level of success in transplants to 
brackish water occurred when plants 3 cm or less in length, 
originating at stations close to the brackish water, were 
transplanted with their rhizomes intact. 

A distinct potential for vegetative propagation was ob- 
vious throughout our transplant studies. In most cases 
transplants of individual Zostera plants responded favor- 
ably. The only exception occurred with transplants from 
intertidal to subtidal locations. 

A lack of clear correlation was evident between the onset 
of the plants' various phenological phases and temperature. 
However, there was some indication that the onset of 
various phenological stages may be, in part, related to 
moderate salinity ranges of 24 to 27 o/oo. In all instances, 
phenological development started in plants nearer the open 
coast and progressed up the estuary. A tendency of Zostera 
to form ecotypes between stations which differed in their 
seasonal salinity patterns was obvious. However, further 
studies to determine the effects of temperature related 
salinity tolerances may be warranted. McMillan (1956) 
has noted a tendency for some Graminae to adapt towards 
different habitats. In addition Biebl and McRoy (1971) 
have noted that Zostera may exhibit a physiological adap- 
tation to a particular environment. This fact was obvious 
in our investigations because Zostera could be collected in 
distinctly subtidal habitats at depths as great as 6 meters. 
Since there is no possibility of these plants ever being ex- 
posed to the atmosphere, we consider them to be good 
examples of adaptation to a subtidal existence. 



466 Rhodora [Vol. 77 



Our results indicate that Zostera marina, may undergo 
successful ecotypical adaptation toward a particular en- 
vironment. This adaptation is probably more closely re- 
lated to moderate salinities than specific temperatures. 
We also have shown that transplants of Zostera between 
habitats is feasible. 

The mathematical formula employed in this investigation 
proved to be an accurate method of assessing biomass for 
Zostera without harvesting the plants. 



LITERATURE CITED 

BlEBL, R., & C. P. McROY. 1971. Plasmic resistance and rate of 

■ 

respiration and photosynthesis of Zostera marina at different 
salinities and temperatures. Marine Biol. 8: 48-5(3. 

JEPPS, M. W. 1931. Note on a marine Labyrinthula. Jour. Mar. 
Biol. Assoc. U. K. 17: 833-838. 

McRoy, C. P. 1970. Standing stocks and other features of eel- 
grasses (Zostera marina) populations on the coast of Alaska. 
Jour. Fish. Res. Board Canada 27: 1811-1821. 

Renn, C. E. 1934. Wasting disease of Zostera marina in Ameri- 
can waters. Nature 134(3385): 416. 

. 1935. A mycetozoan parasite of Zostera marina. 



Nature 135: 544, 545. 



1936a. The wasting disease of Zostera marina* Biol. 



Bull. 70: 148-158. 



. 1936b. Persistence of the eelgrass disease and para* 

site on the American Atlantic coast. Nature 138: 507-508. 

Young, E. L. 1937. Notes on the labyrinthulan parasite of the eel- 
grass, Zostera marina. Bull. Mt. Desert Island Biol. Lab. pp. 
33-35. 



. 1943. Studies on Labyrinthula, the etiologic agent of 

the wasting disease of eelgrass. Am. Jour. Bot. 30: 586-593. 



DEPARTMENT OF BOTANY AND PLANT PATHOLOGY 

AND JACKSON ESTUARINE LABORATORY 
UNIVERSITY OF NEW HAMPSHIRE 
DURHAM, NEW HAMPSHIRE 03824 



OBSERVATIONS ON 
BATRACHOSPERMUM (RHODOPHYTA) 

IN SOUTHEASTERN WISCONSIN STREAMS 



William J. Woelkerling 



mum 



in Wisconsin stems mainly from the report of Prescott 



(1951) 



from 



without reference to locality and with only scant ecological 
data. Moreover, information on seasonal periodicity and 
environmental conditions of the type obtained by Dillard 
(1966) in North Carolina, Minckley and Tindall (1963) in 
Kentucky, and Rider and Wagner (1972) in Pennsylvania 
apparently is lacking for Wisconsin. (The last reference 
includes a literature review of Batrachospermum ecology.) 
The present investigation has been undertaken to gain 
information on 1) the occurrence of Batrachospermum in 
southeastern Wisconsin streams and 2) the environmental 
conditions present at localities where this taxon grows. 
During the course of the study 201 randomly selected 
stream localities have been visited; Batrachospermum 
plants occurrec. at 13 or 6.4% of these sites. 



MATERIALS AND METHODS 

At each station (Table 1) where Batrachospermum 
plants were found, chemical and physical data on alkalinity, 
carbon dioxide, hardness (calcium and total), nitrate ni- 
trogen, orthophosphate, oxygen, pH, temperature, and 
turbidity have been gathered using a Hach Water Analysis 
Field Kit, model DR-EL, which employs microadaptations 
from the "Standard Methods" handbook (American Public 
Health Association 1965). In addition, some observations 
on the type of substrate, relative current velocity, and 
relative exposure to sunlight have been made. 

At stations where Batrachospermum populations per- 
sisted for extended periods, observations were made at 
4-6 week intervals for 10-11 months to note any seasonal 
changes. 



167 



468 



Rhodora 



[Vol. 77 





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1975] 



Batrachospermum — Woelkerling 



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470 Rhodora [Vol. 77 



Voucher specimens from all localities have been collected 
and immediately preserved in FAA (10:7:2:1 95% etha- 
nol:water:formalin:glacial acetic acid). Dried herbarium 
specimens (bearing numbers prefaced by WJW) and per- 
manent microscope slides using" KARO as a mountant 
(Woelkerling, 1970) as well as liquid preserved material 
have been retained in the author's personal collections, 
currently housed at WIS. Species determinations have 
been made primarily with the aid of the taxonomic key of 
Tsraelson (1942); the papers of Kylin (1912), Prescott 
(1951), Sirodot (1884), and Whitford and Schumacher 
(1969) also have been consulted. 



RESULTS AND DISCUSSION 



mum 



have been identified to species. Batrachospermiim borya- 
num Sirodot, not recorded previously from Wisconsin, 
occurred at seven localities (B, F, K, SC, SR, SS, T; ab- 
breviations explained in Table 1), and with one exception 
(locality B), it always grew mixed with other Batracho- 



spermum 



from 



4-21 cm with most plants averaging about 8-10 cm call. 



months 



Ma 



show that sexual plants do occur throughout the year. 

Batrachospermum moniliforme Roth has been found at 
ten localities (BE, F, J, K, MC, N, SC, SR, SS, T), and 
has been reported previously from Wisconsin (Prescott, 
1951). Except for two stations (J, N), it grew mixed with 
other Batrachospermum taxa. Sexually mature plants oc- 
curred throughout the year and ranged in length from 
3-25 cm with an average height of 6-9 cm. 

In addition to the above two taxa, sterile (and thus spe- 
cifically unidentifiable) plants of Batrachospermum have 
been encountered at five localities (BE, M, Q, SC, T) from 
January-March, May-July, and in December. They varied 
in length from 3 cm to 15 cm. The size of some Wisconsin 



1975 1 Batrachospermum — Woelkerling 471 



plants (up to 25 cm) greatly exceeds the 6 cm maximum 
recorded by Rider and Wagner (1972) and the 10 cm maxi- 
mum found by Israelson (1942). 

All Wisconsin populations of Batrachospermum observed 
during this investigation grew at or near the headwaters 
of spring-fed streams, thus agreeing with the findings of 
Minckley and Tindall (1963). Rider and Wagner (1972) 
also recorded their taxa from a spring-fed stream but 
without mention of the headwater areas. Current veloci- 



Wi 



10 cm /sec 



and in most cases exceeded 25 cm /sec. In addition, all 
Wisconsin localities but one (BE) contained rocky or 



>m 



siltation and high turbidity levels. 

Except for seasonal fluctuations in temperature and 
diurnal fluctuations in carbon dioxide and oxygen levels, 
chemical ana physical conditions at any one locality tended 
to remain within relatively narrow limits during the study 
period. Conditions did vary considerably between localities, 
however, and the taxa of Batrachospermum encountered 
appear to tolerate a fairly wide range of environmental 
conditions (Table 2). 



Of particular note is the variation in carbon dioxide 
levels. At no time did CO, levels exceed 24 ppm, and levels 
as low as 1 ppm have been encountered. These values are 
decidedly lower than those reported by Minckley and Tin- 
dall (1963) for Batrachospermum sp. and by Rider and 
Wagner (1972) for B. vagum, but they more or less agree 
with the range in values measured by Rider and Wagner 
(op. cit.) for B. moniliforme. Since both species found in 
Wisconsin (B. moniliforme and B. boryanum) apparently 
require free CO. for photosynthesis (Ruttner, 1960), data 
from the present study strongly suggest that these taxa 
can survive at very low concentrations of free CO., at 
least for short periods of time. 

At six of the thirteen localities (B, J, M, MC, N, Q), 
Batrachospermum plants were encountered on only one 
occasion (Table 1), and data for three additional localities 



472 



Rhodora 



[Vol. 77 



Table 2 

RANGE IN ENVIRONMENTAL CONDITIONS 
UNDER WHICH BATRACHOSPERMUM OCCURRED 



B. boryanum B. moniliformis B. spp. 



15-342 ppm 



2-23 ppm 



2-340 ppm 



Alkalinity 
Carbon 

Dioxide 
Hardness, 

Calcium 
Hardness, 

Total 
Nitrate 
Oxygen 

pH 

Ortho- 
phosphate 0.007-4.0 ppm 
Temperature 6°-18°C 



22-442 ppm 
1.0-4.8 ppm 
8-20 ppm 
6.2-8.4 



Turbidity 



0-7 J.U. 



12-387 ppm 



1-23 ppm 



15-410 ppm 



29-500 ppm 

0.2-16.5 ppm 

5-21 ppm 

6.7-8.4 



02-4.9 ppm 

1°-21°C 
0-10 J.U. 



287-368 ppm 



2-20 ppm 



176-408 ppm 



352-420 ppm 
2.0-10.0 ppm 
5-17 ppm 

7.2-8.2 



02-8.1 ppm 
4°-22°C 
0-15 J.U. 



Table 3 

RANGE IN SELECTED ENVIRONMENTAL 
CONDITIONS AT SCUPPERNONG CK. (SC) 



TICHIGAN 



(T) 



STUDY 



Environmental Factor 



SC 



T 



Alkalinity 
Carbon Dioxide 
Hardness, Ca 
Hardness, Total 

PH 

Temperature 



275-310 ppm 

2-23 
190-225 ppm 

350-400 ppm 

7.5-8.4 

4-15°C 



) ppm 

ppm 

)ppm 



7.7-8.3 



ppm 



5-22°C 



1975] Batrachospermum — Woelkerling 473 



(BE, SR, SS) are confined to three or four dates. Conse- 
quently, information on seasonal changes is restricted to 
limited observations at four localities (F, K, SC, T) from 
either May or June, 1972 through April, 1973. Batracho- 
spermum boryanum and B. moniliforme occurred in mixed 
populations at all four sites; consequently, reference to 

n in the ensuing discussion includes both 



rmu 



taxa. 

The Fontana population (F) grew in very hard water 

(Ca hardness^ 250 ppm CaCO,; total hardness 410 ppm 
CaCO ; ), swift flowing stream 1-2 m across whose tem- 
tures ranged from 12°C in summer to 6°C in winter. Car- 
bon dioxide levels varied from 2-23 ppm, pH from 7.8-8.4, 
and alkalinity from 330-390 ppm. The stream bottom was 
primary gravel. During most of the day the habitat was 

exposed to full sunlight. 

When first discovered in June, 1972, Batrachospermum 
plants occurred in considerable numbers and reached 
lengths of up to 18 cm. By July, 1972, however, most of 



the plants had disappeared or were obviously m 
Several plants (preserved as WJW 3936) appeared heavily 
calcified. The population disappeared entirely by August, 
and new adult gametophytes did not become apparent until 



cm 



were discovered. By January, 1973, Batrachospermum had 
become the dominant alga in the stream with most plants 
averaging 3-6 cm in length. Throughout the remainder of 
the study period (ending in April, 1973), Batrachosper- 



maintained its 



im 



cm 



Similar seasonal fluctuation in population levels also oc- 
curred at Koshwego Springs (K), where certain chemical 
and physical conditions differed considerably from those 
at Fontana. At Koshwego, the water was very soft (Ca 
hardness ^ 15 ppm CaCO.; total hardness ~ 25-30 ppm 
CaCO,), acid (pH varied from 6.2-7.0), and showed alka- 
linity readings of 12-34 ppm. Stream bottom varied from 
rocky to sandy to partially silty with Batrachospermum 



474 



Rhodora C Vo1 - 77 



confined to rocky areas. At all times, the habitat was sub- 
jected to deep shade. In other respects the two localities 
appear more or less similar; at Koshwego, temperature 



from 12°C in summer 



ppm 



May 



mum 



earn vecretation, but bv mi 



become very moribund or had disappeared. Small (i.e., 
less than 2.5 cm tall) plants reappeared in considerable 
numbers in November, 1972, and dominated the stream 
vegetation throughout the winter. Noticeable increase in 
size occurred between February (average size under 2.5 
cm) and March, 1973 (average size 6 cm). Severe flooding 
and silting of the stream occurred in late March and early 
April, 1973, and the Batrachospermum population was al- 
most entirely destroyed. 

At the remaining two stations (SC, T) Batrachospermum 
plants occurred throughout the year and formed the domi- 
nant component of the algal vegetation during much of that 
time. Both stations had environments (Table 3) similar to 
that at Fontana except that one (SC) was largely shaded 
throughout the day and the other (T) was exposed to full 
sunlight during most of the day. In addition the latter (T) 
had summer temperatures of 18-22°C or 5-10°C higher 
than at the other hard water localities. 

Immediately below the spring from which Scuppernong 
Creek originates, Batrachospermum plants constituted the 
dominant form of vegetation. Within 100 m, however, 
angiosperm vegetation became dominant and the Batracho- 
spermum- population consisted only of scattered plants. 
During winter months most plants encountered were 4-6 cm 
long and during summer they were 8-10 cm long; one 15 
cm tall plant was encountered. 

The Tichigan Creek population of Batrachospermum 
dominated the macroscopic vegetation throughout the year. 
Summer plants generally did not exceed 10 cm in length; 
winter plants all (i.e., December- April) were enormous in 
size and reached lengths of up to 25 cm. The very large 



1075] Batrachospermum — Woelkerling 475 



size of these individuals as compared to the other winter 
populations studied could not be accounted for on the basis 
of the physical or chemical parameters examined during 

the study. 

The above observations suggest that different populations 

of Batrachospermum (at least in Wisconsin streams) may 

either produce mature gametophytes throughout the year 

(SC, T) or show seasonal variation with an absence of 



m 



mid-summer 



suggest that maximum vegetative development can occur 
in spring (F, K), summer (SC), or winter (T). Previous 

can studies (Dillard, 1966, Minckley and Tindall, 



m n 



American studies (Dillard, 1966, 
1968, Rider and Wagner, 1972) all reported definite sea- 
sonal fluctuations in Batrachospermum populations with a 
disappearance of plants in summer and a reappearance in 
fall. Yoshida (1959), however, makes mention of both 
seasonal and year-round populatioi 
in Japanese streams. 

Various attempts have been made to account for seasonal 
fluctuation in population levels in terms of temperature 
changes, changes in light intensity, and differences in cur- 
rent velocity (see Dillard, 1966, Minckley and Tindall, 
1963, Rider and Wagner, 1972, Yoshida, 1959). The re- 
sults of the Wisconsin study, however, indicate that mature 
plants and maximum vegetative development can occur 
under both low and high light intensities and under both 
summer and winter temperatures. Therefore, other fac- 
tors, perhaps genetic, appear to be involved in determining 
why different populations of the same species either persist 
year round or show seasonal fluctuations. No relationship 
to current velocity has been observed in this investigation. 

The apparently consistent occurrence of Batrachosper- 
mum in the headwater areas of spring-fed streams likewise 
requires further investigation. Minckley and Tindall (1963) 
suggest that the availability of unbound carbon dioxide 
may be a controlling factor (their stream reportedly has 
super-saturated CO, levels), but the relatively low CO, 
levels found during this study again suggest that other 



476 



Rhodora [Vol. 77 



factors may be involved, and additional study appears 
warranted. 



SUMMARY 



some 



spennum in southeastern Wisconsin streams have been 



num 



Wisconsin), B. monilif 



occurred in 6.4% of the localities visited and were found 
in both alkaline, hard water and acid, soft water environ- 
ments. Depending upon the population, mature plants per- 
sisted throughout the year or disappeared in summer and 
fall, and they showed maximum vegetative development in 

or winter. The seasonal behavior does 



summer 



not appear to be correlated entirely with changes in light 
intensity or temperature. Likewise the apparent occurrence 
of Batrachospermum near the headwaters of spring-fed 
streams apparently cannot be explained solely on the basis 
of greater availability of unbound carbon dioxide in these 
habitats. 



ACKNOWLEDGMENTS 



Sincere thanks are due Mr. Robert Dietrich and Mr. 
Warren Mueller for assistance in the gathering and pro- 
cessing of data. This study was supported by grant No. 
130376 from the Research Committee of the University of 
Wisconsin Graduate School. 



LITERATURE CITED 

American Public Health Association, The American Water Works 

Association, and the Water Pollution Control Federation. 1965. 
Standard Methods for the Examination of Water and Waste- 
water, ed. 12, 769 pp. New York. 
DlLLAED, G. E. 1966. The seasonal periodicity of Batrachospermum 
macroaporum and Audouinella violacea in Turkey Creek, North 
Carolina. Jour. Elisha Mitchell Sci. Soc. 82: 204-207. 

Israelson, G. 1942. The freshwater Florideae of Sweden. Symh. 
Bot. Upsal. 6: 1-135. 



1975] Batrachospermum — Woelkerling 477 



Kylin, H. 1912. Uber die roten und blaue Farbstoffe der Algen. 
Hoppe-Seylers Z. Physiol. Chem. 76: 397-42.1. 

MlNCKLEY, W. L., & D. R. Tindall. 1963. Ecology of Batracho- 
spermum sp. in Doe Run, Meade County, Kentucky. Bull. Tor- 
rey Rot. Club 90: 391-400. 

PRESCOTT, G. W. 1951. Algae of the Western Great Lakes Area. 

946 pp. Cranbrook Inst. Sci. 

Riper, D. E„ & R. H. Wagner. 1972. The relationship of light, 
temperature, and current to the seasonal distribution of Batra- 
chospermum (Rhodophyta). Jour. Phycol. 8: 323-331. 

Ruttner, F. 1960. Carbon uptake in algae of Rhodophycean genus 

Batrachospermum. Schweiz. Z. Hydrol. 22: 280-291. 

Sirodot, S. 1884. Les Batrachospermes. Organisation, functions, 
developement, classification. 299 pp. Paris. 

Whitforl), L. H., & G. J. Schumacher. 1969. A Manual of the 

Freshwater Algae in North Carolina. 313 pp. Raleigh, N. C 

Woelkerling, W. J. 1970. Acrochaetium botryocarpum (Harv.) 

J. Ag. (Rhodophyta) in southern Australia. Br. Phycol. Jour. 

5: 159-171. 
YOSHIDA, T. 1959. Life cycle of a species of Batrachospermum 
found in Northern Kyushu, Japan. Jap. Jour. Bot. 17: 29-42. 



Department of Botany 
University of Wisconsin 
Madison, Wisconsin 53706 



A COUNTY CHECKLIST OF THE FERNS 

AND FERN ALLIES OF KANSAS, NEBRASKA, 

SOUTH DAKOTA, AND NORTH DAKOTA 1 



Aleta Jo Petri k-Ott 



INTRODUCTION 

To the present, there has been no comprehensive work 
dealing- exclusively with the ferns and fern allies of what 
is considered to be the heart of the central plains and 
prairies of the United States, i.e., Kansas, Nebraska, South 
Dakota, and North Dakota. 

Of the larger floras, Rydberg's "Flora of the Rocky 
Mountains and Adjacent Plains" (1917) extends eastward 
to longitude 102° W in Nebraska, South Dakota, and North 
Dakota. Rydberg's "Flora of the Prairies and Plains of 
Central North America" (1932) includes the ferns and fern 
allies of the aforementioned four states, but several taxa 
reported to be in these states have never been verified, 
many new taxa have been found since that time, and recent 
research has resulted in many nomenclatural changes. 
Fernald's account of the ferns and fern allies in "Gray's 
Manual of Botany" (1950) covers only the area in Ne- 
braska and Kansas east of longitude 96° W and thus 
excludes the prairie and plains regions of the Dakotas, 
Nebraska, and Kansas. 

Smaller, individual state floristic treatments briefly cov- 
ering the ferns and/or fern allies are represented by Keller- 
man and Kellerman (1888), Saunders (1899), Bergman 
(1918), Petersen (1923), Over (1932), Gates (1940), 
Stevens (1950), and Winter, Winter, and Van Bruggen 
(1959). Floristic regional and state studies exclusively 
concerning ferns and/or fern allies are represented by 
Wilson (1885), Cragin (1885, 1886), Bessey (1892), Reed 
(1896), Fitzpatrick (1919, 1920a & b), Humfeld (1951a 

■Contribution Number 554, Virginia Institute of Marine Science, 
Gloucester Point, Virginia 23062. 



478 



1975] Ferns — Petrik-Ott 479 



McGregor 



McGregor (1960), 



Brooks (1967, 1969), and Van Bruggen (1967). 

Supporting specimens are not to be found for many 
records given in the existing literature, and many of the 
older records were based upon mis-identifications. The 
rwnrrU o-ivpn herp are based entirely upon specimens from 



mai 



tions which are deposited in the University of Kansas 

The nine herbaria are (abbreviations after 



arium 



Lanjouw and Stafleu, 1964) : 

kanu = The University of Kansas, Lawrence, Kansas 



KSC 



Manhattan 



KSTC = Herbarium of the Kansas State Teachers 

College, Emporia, Kansas 
MO = Missouri Botanical Garden, 2815 Tower 

Grove Avenue, Saint Louis 10, Missouri 
NDA = North Dakota State University and Experi- 



ment Station, Fargo, North 



NEB 



Museum 



coin 8, Nebraska 



SDC = Department of Botany, South Dakota State 

College, College Station, Brookings, South 
Dakota 

SDU = University Herbarium, University of South 

Dakota, Vermillion, South Dakota 

umo = University of Missouri Herbarium, Colum- 
bia, Missouri 

The following list was compiled during the years 1968- 
1970. Citations for each taxon are grouped alphabetically 
by state and county. Only one specimen for each county 
was chosen as representative. A map (Fig. 1) showing the 
counties in each state is included. To conserve space, it was 
considered sufficient to give only the collector's name and 
his collection number for each representative county speci- 



men 



numb 



arium 



480 



Rhodora [Vol. 77 



if indicated, was designated. It is realized that the county 
distributions of certain taxa have probably been extended 
since the termination of my study, but it has not come to 
my attention that any new taxa have been added. 

Because there has been much confusion concerning the 
presence of certain taxa, which cannot be verified by exist- 
ing specimens, a section on doubtful collections and ex- 
cluded taxa has been appended to the checklist. 



THE FERNS 



Family Ophioglossaceae 



Botrychium dissectum Sprang, var. dissectum Anleit. 
Kennt. Gewachse 3:172. 1804. 

Kansas: Cherokee CO.: R. L. McGregor 3870 (kanu). 

Botrychium dissectum Spreng. var. obliquum (Muhl.) 
Clute, Fern Bull. 10:76. 1902. 

Kansas: Anderson Co.: L. K. Mag rath 4988 (kanu). bourbon 
CO.: S. Stephens 19492 (kanu). Cherokee CO.: R. L. McGregor 
3860a (kanu). coffey co.: L. K. Maffrath 4979 (kanu). douglas 
CO.: J. E. Bare 762 (kanu). franklin CO.: L. K. Magrath 5086 
and A. Organ (k\nu). Jefferson CO.: R. L. McGregor 4154 
(kanu). linn CO.: L. K. Magrath 4938 (kanu). miami CO.: W. H. 
Horr and R. L. McGregor 3617 (kanu). wyandotte Co.: L. K. 
Magrath 5056 (kanu). 

Botrychium lunaria (L.) Swartz, Schrad. Journ. Bot. 1800 
(2):110. 1801. 

North Dakota: mchenry CO.: O. A. Stevens 1530 (nda). 

Botrychium multifidum (S. G. Gmel.) Rupr. Beitr. Pflan- 
zenk. Russ. Reich. 11:40. 1859. 

Nebraska: franklin CO.: E. M. Hiissong 4fi89 (neb). 

South Dakota: CUSTER CO.: S. Stephens 35426 and R. Brooks 

(kanu). pennington CO.: T. Van Brag gen 5042 (sdu). 

Botrychium simplex E. Hitchc. Amer. Journ. Sci. 6:103. 
pi. 8. 1823. 

South Dakota: Lawrence Co.: C. A. Taylor 7963 (sdc). 



1975] 



Ferns 



Petrik-Ott 



1S1 




Figure 1. County map of Kansas, Nebraska, South Dakota, and 
North Dakota. 



482 Rhodora [Vol. 77 



Botrychium virginianum (L.) Swartz, Schrad. Journ. Bot. 
1800(2) :111. 1801. 

Kansas: Anderson CO.: R. L. McGregor 3211 (kanu). atchison 
CO.: A. J. Petrik-Ott 752 (kanu). bourbon CO.: A. J. Petrik-Ott 
756 (kanu). brown eo.: R. L. McGregor 2836 (kanu). chase Co.: 
B. B. Smyth, 19 Aug-., 1902 (ksc). Chautauqua co.: P. H. Hum f eld 
842 (kanu). Cherokee CO.: A. J. Petrik-Ott 724 (kanu). COFFEY 
CO.: L. K. Magrath 3876 (kstc). cowley CO.: W. H. Horr and 
R. L. McGregor, 23 July 1947 (kanu). Crawford co. : A. J. Petrik- 
Ott 725 (kanu). doniphan CO.: R. L. McGregor 12258 (kanu). 
douglas CO.: R. L McGregor 14217 (kanu). franklin CO.: R. L. 
McGregor 5592 (kanu). geary CO.: A. £. 'Hitchcock, June 1896 
(ksc). greenwood CO.: S. Stephens 2914 (kanu). jackson CO.: 
R. L. McGregor 2853 (kanu). JEFFERSON CO.: R. L. McGregor 880 

(kanu). johnson co.: P. H. Hum f eld 595 (kanu). Leavenworth 
CO„: P. H. Humfeld 195 (kanu). linn co. : A. J. Petrik-Ott 763 
(kanu). lyon co.: J. S. Wilson 2042 (kstc). Marshall co.: S. 

Stephens 3291 (kanu). MIAMI CO.: S. Stephens 2312 (kanu). 
Montgomery CO.: R. L. McGregor 12824 (kanu). nemaha CO.: 
L. K. Magrath 4506 (kanu). OSAGE CO.: S. Stephens 30602 (kanu). 
Pottawatomie CO.: S. Stephens 4710 (kanu). RILEY CO.: Miss 
Thackrey, 24 May 1892 (ksc). saline CO.: J. Hancin 375 (ksc). 
stiawnee CO.: L. D. Voile 715 (kanu). WABAUNSEE Co.: S. Stephens 
4798 (kanu). wilson co. : L. C. Hulbert 3686 (ksc). woodson co. : 
R. L. McGregor 905 (kanu). Wyandotte co. : A. J. Petrik-Ott 747 

(KANU). 

Nebraska: BURT CO.: S. Stephens 32072 and R. Brooks (kanu). 
cass CO.: A. J. Petrik-Ott 776 (kanu). cherry CO. : /. M. Bates 

6043 (neb). Dakota CO.: S. Stephen 32048 and R. Brooks (KANU). 

dawes CO.: H. J. Webber 6103 (neb), douglas co.: W. Cleburne, 
9 July 1897 (neb), gage co.: H. L. Shantz, June 1904 (neb), hall 

CO.: J. M. Bates 3807 (neb), holt CO. : R. Brooks 374 and S. Stephens 

(kanu). Lancaster co.: H. J. Webber 6102 (neb), nemaha co.: 
S. Stephens 3660 (kanu). otoe CO.: H. J. Webber, 1 June 1889 
(mo), rici/ardson co.: A. J. Petrik-Ott 773 (kanu). sarpy co.: 
W. Cleburne, 3 June 1887 (neb). SIOUX Co.: A. F. Woods 444 (neb), 
thomas CO.: P. A. Rydberg 1467 (neb). THURSTON Co.: S. Stephens 

21 066 (kanu). 

North Dakota: BENSON CO.: H. F. Bergman 1890 (nda). BOTTI- 
NEAU CO.: H. F. Bergman 2550 (nda). CASS CO.: H. F. Bergman 
1637 (nda). dunn CO.: O. A. Stevens and D. R. Moir, 14 Sept. 
1956 (nda). pembina CO.: L. R. Waldron 1649 (nda). ransom CO.: 
S. Stephens 33426 and R. Brooks (kanu). richland CO.: O. A. 
Stevens 424 (nda). stark CO.: Mrs. M. Meissncr, 20 Au£. 1939 
(nda). 



1975] Ferns — Petrik-Ott 48 






South Dakota: clay CO.: G. B. Smith, 22 Sept. 19(5(5 (sdu). Hard- 
ing co.: S. S. Visiter 7057 (sdu). lawrence CO.: S. Stephens 7395 
(kanu). Lincoln CO.: N. Winter, Oct. 1954 (sdu). Marshall CO.: 
T. Van Brvgggen 4448 (sdu). Roberts Co.: R. Brooks 443 (kanu). 

Ophioglossum engelmanni Prantl, Ber. Deutsch. Bot. Ges. 
1:351. 1883. 

Kansas: ALLEN CO.: L. K. Magrath 4977 (kanu). ANDERSON co. : 
P. H. Thompson, 19 June 1949 (kanu). ATCHISON CO.: P. L. Mc- 
Gregor 887 (kanu). bourbon CO.: S. Step/tens 3978 (kanu). Chau- 
tauqua co.: P. L. McGregor 4186 (kanu). Cherokee CO.: R. L. 
McGregor 15328 (kanu). COFFEY CO.: L. K. Magrath 3836 (kstc). 
COWLEY CO.: Rudy G. Koch 3468 (KSC). DOUGLAS CO.: P. H. Hum f eld 

1011 (kanu). elk co.: S. Stephens 10754 (kanu). johnson co.: 

R. H. Thompson, 22 April 1941 (kanu). LEAVENWORTH CO.: R. L. 

McGregor 886 (kanu). miami co.: P. H. Humfeld 219 (kanu). 

MONTGOMERY CO.: P. H. Hum 1 eld 107 (kanu). neosho CO.: W. W. 
Holland, 2622 (kanu). wilson CO.: R. L. McGregor, 30 June 1917 

(kanu). 

Ophioslossum vulgatum L Sp. PL 2:1062. 1753. 

Nebraska: cherry CO.: ,/. M. Bates 5679 (neb). 



Family Osmundaceae 

Osmunda regal is L. var. speetabilis (Willd.) A. Gray, Man. 
ed. 2:600. 1856. 

Kansas: wilson CO.: L. L. Kiefer, 5 Oct. 1961 (kanu). woodson 
cc- W. H. Horr y 10 July 1930 (kanu). 
Nebraska: E. 71/. Hussong 6776 (neb). 



Family Adiantaceae 

Adiantum capillus-veneris L. Sp. PI. 2:1096. 1753. 

South Dakota: fall river CO.: A. J. Petrik-Ott 784 (kanu). 

Adiantum pedatum L. Sp. PL 2:1095. 1753. 

Kansas: allen CO.: R. L. McGregor 968 (kanu). atchison CO.: 
P. L. McGregor 2814 (kanu). bourbon CO.: R. L. McGregor 979 

(KANU). BROWN CO.: P. L. McGregor 904 (KANU). CHEROKEE CO.: 

R. L. McGregor 983 (kanu). coffey CO.: P. L. McGregor 918 

(KANU). CRAWFORD CO.: P. L. McGrogcr 981 (kanu). DONIPHAN 

CO.: A. ./. Petrik-Ott 753 (kanu). DOUGLAS CO.: P. L. McGregor 
17356 (kanu). franklin CO.: P. L. McGregor 10043 (kanu). 
jackson co.: M. Reed, 1 Sept. 1889 (ksc). Jefferson co.: P. L. 



484 Rhodora [Vol. 77 



McGregor 885 (kanu). Johnson co.: P. H. Humfeld 611 (kanu). 
Leavenworth CO.: R. L. McGregor 2792 (kanu). linn CO.: A. J. 
Petrik-Ott 762 (kanu). Miami CO.: R. L. McGregor 896 (kanu). 
neosho CO.: M. D. Eiving, Summer 1891 (ksc). osage CO.: R. L. 
McGregor 891 (kanu). shawnee Co.: L. D. Voile 131 (kanu). 
wtlson Co.: H. Danell, June 1890 (ksc). woodson CO.: R. L. Mc- 
Gregor 917 (KANU). WYANDOTTE CO.: 5. Stephens 10863 (kanu). 
Nebraska: cass CO.: S. Stephens 3743 (kanu). cuming CO.: L. 
Brunrr, July 1880 (neb), douglas CO.: W. Cleburne, June 1873 
(neb). Lancaster CO.: H. J. Webber 6128 (neb), nemaha CO.: 
J. M. Bate? 5145 (neb), otoe CO.: K. L. Johnson 1584 and D. E. 

Dallas (kanu). pawnee CO.: J. E. Shue, May 1896 (neb). Richard- 
son CO.: A, J. Petrik-Otl 770 (kanu). rock co. : J. M. Bates, 
13 Aug 1 . 1900 (neb), sarpy CO.: W. L. Tolstead and H. C. Reynolds, 
1.7 May 1942 (neb). 

Cheilanthes alabamensis (Buckley) Kunze, Linnaea 20:4. 
1847. 

Kansas: Cherokee co.: R. L. McGregor 3865 (kanu). 

Cheilanthes feei Moore, Index Fil. 38. 1857. 

Kansas: barber CO.: J?. L. McGregor 14950 (kanu). Chautauqua 
CO.: A. J. Petrik-Ott 745 (kanu). Ellsworth CO.: R. L. McGregor 
1312 (kanu). ford CO.: R. L. McGregor 10928 (kanu). hodgeman 
CO.: R. L. McGregor 10935 (kanu). Lincoln co. : D. E. Lantz, 
3 May 1898 (ksc). Ottawa co.: A. J. Petrik-Ott 769 (kanu). 
russell CO.: L. C. Hulbert 4346 (ksc). stanton co.: R. L. Mc- 
Gregor 16115 (kanu). 

Nebraska: banner co. : P. A. Rydberg 479 (neb), scotts bluff 
CO.: S. Stephens 5481 (kanu). 

South Dakota: custer CO.: S. Stephens 6167 (kanu). fall river 
CO.: S. Stephens 5797 (kanu). harding CO.: S. Stephens and R. 
Brooks 425 (kanu). Lawrence co. : S. Stephens 7475 (kanu). 
meade CO.: E. J. Palmer 37017 (mo), pennington Co.: S. Stephens 

7134 (KANU). 

Cheilanthes lanosa (Michx.) D. C. Eat. in Torr. Rep. U. S. 



Mex 



1859. 



Kansas: Chautauqua co. : A. J. Petrik-Ott 7 '45 (kanu). Cherokee 
CO.: R. L. McGregor 11063 (kanu). elk co.: R. L. McGregor 926 
(kanu). Montgomery CO.: R. L. McGregor 946 (kanu). WILSON 
CO.: R. L. McGregor 925 (kanu). woodson Co.: R. L. McGregor 
914 (kanu). 

Cheilanthes tomentosa Link, Hort. Berol. 2:42. 1833. 

Kansas: Cherokee CO.: R. L. McGregor 3866 (kanu). 



1975] Ferns — Petrik-Ott 485 






Notholaena dealbata (Pursh) Kunze, Amer. Journ. Sci. II. 
6:82. 1848. 

Kansas: allen co.: R. L. McGregor 3335 (kanu). ANDERSON 
CO.: R. L. McGregor 3209 (kanu). bourbon CO.: A. J. Petrik-Ott 
759 (kanu). Chautauqua co. : R. L. McGregor 951 (kanu). Chero- 
kee CO.: R. L. McGregor 971 (kanu). COFFEY CO.: R. L. McGregor 

910 (kanu). cowley co.: R. G. Koch 1764 (kanu). Crawford co. : 

R. L. McGregor 973 (kanu). DOUGLAS Co.: R. L. McGregor 874 
(kanu). elk CO.: R. L. McGregor 15880 (kanu). franklin Co.: 
R. L. McGregor 3309 (kanu). geary CO.: C. C. Parry, April 1873 
(mo), greenwood co.: R. L. McGregor 994 (kanu). jefferson co. : 
P. H. Humfeld 132 (kanu). johnson CO.: R. L. McGregor 343.") 

(KANU). LABETTE CO.: R. L. McGregor 955 (kanu). LEAVENWORTH 

CO.: P. H. Humfeld 1019 (kanu). Lincoln CO.: D. E. Lantz, 3 May 
1898 (ksc). linn co.: M. Campbell, 28 June 1890 (KSC). miami 

co.: P. H. Humfeld 214 (kanu). Montgomery CO.: A. ,/. Petrik-Ott 
735 (kanu). morris CO.: C. C. Parry, 29 April 1873 (mo), neosho 
o.: R. L. McGregor 957 (kanu). Pottawatomie CO.: P. H. Hum- 
feld 396 (kanu). riley CO.: W. A. Kellerman 19 Aug". 1887 (ksc). 
shawnee co.: P. B. Smyth 70 (ksc). WILSON co.: />. L. Wagen- 
knecht 2231 (kanu). WOODSON CO.: R. L. McGregor 992 (kanu). 

wyandotte co.: S. Stephens 10858 (kanu). 

Nebraska: CASS co. : T. A. Williams 6773 (neb). 

Pellaea atropurpurea (L.) Link, Fil. Sp. Hort. Reg. Bot. 
Berol. Cul. 59. 1841. 

Kansas: \llen CO.: R. L. McGregor 3332 (kanu). Anderson CO.: 

R. L. McGregor 3209 (KANU). ATCHISON CO.: R. L. McGregor 2811 

(kanu). barber CO.: h\ L. Richards 3716 (kanu). barton co. : 
S. Stephens 8380 (kanu). bourbon CO.: A. J. Petrik-Ott 757 (kanu). 
chautauqua CO.: P. H. Humfeld 855 (kanu). Cherokee CO.: E. L. 

Richards 3104 (kanu). clay co. : R. L. McGregor 5016 (kanu). 

cloud co.: S. V. Fraser 558 (ksc). coffey co. : R. L. McGregor 
913 (kanu). comanche CO.: R. L. McGregor 10944 (kanu). cowley 
co.: 5. Stephens 3020 (kanu). Crawford CO.: C. L. Merritt, Sum- 
mer 1941 (ksc). donipiian co.: F\ Agrelius, Aug. 1913 (kstc). 
Douglas co.: T?. L. McGregor 640 (kanu). elk co.: R. L. McGregor 
15877 (kanu). Ellsworth co.: .R. L. McGregor 17326 (kanu). 
franklin Co.: /?. L. McGregor 889 (kanu). greenwood CO.: S. 
Stephens 9907 (kanu). jefferson co.: P. //. Humfeld 133 (kanu). 

JOHNSON CO.: R. L. McGregor 3871 (kanu). KIOWA CO.: .4. S. 

Hitchcock, Aug. 1896 (ksc). Leavenworth co.: R. L. McGregor 

28C6 (kanu). Lincoln co.: P. H. Ham f eld 965 (kanu). LYON CO.: 

F. Agrelius, 10 April 1946 (kstc). Marshall co.: S. Stephens 4666 
(kanu). mcpherson co,: /?. L. McGregor 13720 (kanu). Miami 



486 



Rhodora [Vol. 77 



CO.: P. H. Hamfeld 160 (kanu). Montgomery co. : A. J. Petrik-Ott 
736 (kanu). neosho CO.: W. W. Holland 872 (kanij). osage CO.: 
R. L. McGregor 3318 (kanu). Ottawa CO.: A. J. Petrik-Ott 768 
(kanu). rice CO.: R. L. McGregor 2706 (kanu). riley Co.: D. J. 
Cashen, 24 Oct. 1920 (ksc). RUSSELL CO.: R. L. McGregor 12676 
(kanu). saline CO.: R. L. McGregor 2689 (kanu). shawnee CO.: 
P. H. Hamfeld 145 (kanu). wabaunsee CO.: /?. R. Smyth, 8 Auu*. 
1890 (umo). wilson CO.: B. L. Wagenknecht 2217 (kanu). wood- 
son CO.: /?. L. McGregor 915 (kanu). WYANDOTTE CO.: S. Stephens 

10860 (KANU), 

Nebraska: box butte CO.: M. P. Somcr, Au#. 1909 (neb). Cass 
CO.: H. J. Webber, 23 May 1887 (neb), jefferson co. : A. F. Woods 

and D. Saunders 2001 (neb), nemaha co.: J. M. Bates 6683 (neb). 
South Dakota: custer CO.: S. Stephens 6158 (kanu). fall river 
co.: S. Stephens r>796 (kanu). lawrence co. : C. A. Taylor 7464 
(sdc). meade co.: E. J. Palmer 37019 (mo), pennington co.: 
S. Stephens 7133 (kanu). 

Pellaea glabella Mett. ex Kuhn, Lirmaea 36:87. 1869. 

Kansas: ALLEN CO.: R. L. McGregor 3331 (kanu). bourbon CO.: 

R. L. McGregor 3340 (kanu). chase CO.: F. Agrelius, 9 Aug-. 1953 
(kstc). cuautauqua co.: P. H. Hum f eld 846 (kanu). cowley CO.: 
S. Stephens 3020 (kanu). douolas CO.: R. L. McGregor 649 (kanu). 

franklin CO.: /?. L. McGregor 892 (KANU). GEARY CO.: P. H. Hum- 

feld 967 (kanu). greenwood co. : P. H. Hum f eld 799 (kanu). John- 
son co.: /?. /",. McGregor 3433 (kanu). labette co.: P. M. Afaua, 
26 May 1927 (ksc). Leavenworth co.: P. H. Hum feld 187 (kanu). 
Lincoln co.: D. E. Lctntz, 3 May 1898 (ksc). linn CO.: P. H. Ham- 
feld 40 (kanu). lyon CO.: F. Agrelius, 10 May 1945 (kstc). mar- 
shall co.: W. H. Horr 4616 (kanu). Montgomery co. : R. L. Mc- 
Gregor 10839 (kanu). Ottawa co.: A. J. Petrik-Ott 767 (kanu). 
Pottawatomie CO.: P. H. Humfeld 395 (kanu). riley co.: W. A. 
Ketterman, 28 May 1887 (ksc). scott co. : F. Agrelius, 16 Aug". 
1912 (kstc). wabaunsee co.: R. L. McGregor 12371 (kanu). wil- 
son co.: P. H. Hamfeld 120 (kanu). woodson CO.: W. H. Horr, 
10 July 1930 (kanu). 

North Dakota: bowman CO. : O. A. Steven* and D. R. Moir 2304 
(NDA). dunn CO.: O. A. Stevens, 8 June 1938 (nda). golden valley 
CO.: S. Stephens 23465 and R. Brooks (kanu). grant co.: W. B. Bell 
1395 (ndv). hettinger CO.: Mrs. M. Meissner y 10 Au£. 1938 (nda). 
Mckenzie CO.: O. A. Stevens and D. R. Moir, 14 Sept. 1956 (nda). 
morton co : O. A. Stevens, 17 June 1945 (nda). Oliver co. : O. A. 
Stevens, 7 Aug-. 1938 (nda). 

South Dakota: clay co.: W. H. Over 1680 (SDU). custer co.: 
P. A. Rydhcrg 1191 (neb), harding CO.: Over and Solem 12758 

(SDU). LAWRENCE CO.: S. Stephens 7518 (KANU). PENNINGTON CO.: 

S. Stephens 7350 (kanu). 



1975] Ferns — Petrik-Ott 487 



Family Aspidiaceae 

Athyrium filix-femina (L.) Roth, Tent. Fl. Germ. 3:65. 
1799. 

Nebraska: Adams CO.: J. M. Rates, 12 July 1913 (neb), boyd Co.: 
J. M. Bates 1094 (neb), brown CO.: F. Clements 2938 (neb). 

North Dakota: cavalier co. : O. A. Stevens 2504 (nda). pembin a 
CO.: R. Brooks 449 (kant). richlandco.: O. A. Stevens 149 (nda). 

South Dakota: CUSTER CO.: A. ./. Petrik-Ott 792 (kanu). Law- 
rence Co.: E. J. Palmer 37532 (mo), minnehaha CO.: C. A. Taylor 
7651 (sdc). pennington Co.: S. Stephens 7175 (kanu). 

Athyrium pyenocarpon (Sprengf.) Tidestr. Elys. Marianum 
1:36. 1906. 

Kansas: LEAVENWORTH CO.: J. Wilson 3729 (kanu). WYANDOTTE 
CO.: K. K. Mackenzie 307 (ksc). 

Cystopteris bulbifera (L.) Bernh. Schrad. Neu. Journ. Bot. 
1(2) :10. 1806. 

South Dakota: Roberts CO.: A. J. Petrik-Ott 800 (kanu). 

Cystopteris fragilis (L.) Bernh. Schrad. Neu. Journ. Bot. 
1 (2) :26. pi. 2. fig. 9. 1806. 

Kansas: CRAWFORD Co.: W. W. Holland 963 (KANU). ELLSWORTH 
Co.: R. L. McGregor 17324 (kanu). johnson CO.: M. A, Carleton, 
25 Autf. 1892 (ksc). Montgomery CO.: F. Agrelius, 28 May 1930 
(kstc). rice co.: A'. L. McGregor 2704 (kanu). riley co. : H. F. 
Bergman, 9 Sept. 1910 (nda). saline co. : R. L. McGregor 2(188 
(kanu). WILSON co.: H. Willis 16 (kstc). woodson CO.: R. Brooks 
973 (kstc). 

Nebraska: antelope CO.: N. F. Petersen 1908 (neb), brown co. : 
. E. Bessey, 23 July 1887 (neb), cass CO.: S. Stephens 18922 
(kanu), cherry CO.: W. L. Tolstead, 25 June 1937 (neb), cuming 
CO.: L. Bruner, July 1880 (neb), custer co. : H. J. Webber 6121 
(neb). Dakota co.: S. Stephens 21144 (kanu). dawes co. : S. 

Stephens 24606 and R. Brooks (kanu). dixon CO.: F. Clements 
2556 (neb), douglas co.: W. Cleburne, 28 June 1873 (neb), holt 
CO.: J. M. Bates, 21 Aug*. 1895 (neb), johnson CO.: No collector 
given, 189? (neb), keya pah a CO.: W. Kierur 23692 (mo). KNOX 
CO.: W. T. Barker 2652 (kanu). Lancaster CO.: T. Walton, 20 
May 1886 (neb). Richardson co. : J. M. Winter, July 1931 (sou). 
saunders CO.: T. A. Williams, 24 June 1890 (neb), seward CO.: 
H. ./. Webber 6116 (neb). Sioux CO.: A. J. Petrik-Ott 783 (kanu). 






thomas co.: P. A, Rydberg 1479 (neb), webster co. : J. M. Bates, 
17 April 1905 (neb). 



488 Rhodora [Vol. 77 



North Dakota: barnes CO.: H. F. Bergman 2292 (nda). benson 
CO.: R. Brooks 456 (kanu). billings CO.: R. Brooks 419 (kanu). 
dunn CO.: R. Brooks 395 (kanu). golden valley CO.: £. Stephens 
23453 and R. Brooks (kanu). grant CO.: O. A. Stevens, 10 July 
1964 (nda). mckenzie CO.: R. L. McGregor and J. E. Bare 1153 

(KANU). MCLEAN CO.: R. Brooks 398 (kanu). MERCER CO.: R. 

Brooks 391 (kanu). MORTON CO.: R. Brooks 389 (kanu). pierce 
CO.: O. A. Stevens, 16 July 1939 (nda). ransom CO.: S. Stephens 
33423 and R. Brooks (kanu). sioux CO.: A. J. Petrik-Ott 797 
(kanu). stark CO.: C. H. Waldron, 23 June 1912 (nda). stutsman 
CO.: C. C. Schmidt, 5 July 1897 (nda). ward CO.: R. Brooks 400 
(kanu). Williams co. : O. A. Stevens, 8 Aug. 1915 (nda). 



South Dakota: brule CO.: R. Brooks 461 (kanu). clay CO.: 



W 



Custer CO.: A. J. Petrik-Ott 789 (kanu). 



dewey CO.: S. Stephens 33798 and R. Brooks (kanu). fall river 
CO.: S. Stephens 5795 (kanu). GRANT CO.: W. H. Over 7033 (sou). 
GREGORY CO.: S. Stephens and R. Brooks 375 (kanu). haakon CO.: 
S. Stephens and R. Brooks 379 (kanu). harding CO.: S. Stephens 
7922 (kanu). Lawrence CO.: A. J. Petrik-Ott 795 (kanu). mar- 
shall CO.: 0. A. Stevens and D. R. Moir, 30 May 1957 (nda). 
meade co.: E. J. Palmer 37069 (kanu). Minnehaha CO.: L. J. 
Harms 2689 (kanu). pennington CO.: S. Stephens 7304 (kanu). 

ROBERTS CO.: S. Stephens 21471 (KANU). STANLEY CO.: S. Stephens 
and R. Brooks 377 (kanu). todd CO.: L. Stanley, 24 May 1966 

(stu). union CO.: T. Van Bruggen 4917 (sdu). washabaugh co.: 

J. E. Bare 1227 (kanu). Ziebach CO.: R. Brooks 380 and S. Stephens 

(kanu). 



Cystopteris protrusa (Weath.) Blasdell, Mem. Torrey Bot. 
Club 21(4) :41, 42.pl 3. 1963. 

Kansas: Anderson CO.: R. L. McGregor 3458 (kanu). ATCHISON 
CO.: R. L. McGregor 2817 (kanu). brown CO.: R. L. McGregor 

2937 (kanu). Chautauqua co.: P. H. Hum f eld 868 (kanu). Chero- 
kee co.: S. Stephens 4136 (kanu). doniphan co.: A. J. Petrik-Ott 
755 (kanu). douglas co.: P. H. Hum f eld 383 (kanu). elk CO.: 
S. Stephens 10708 (kanu). franklin CO.: R. L. McGregor 10041 

(KANU). GREENWOOD CO.: R. L. McGregor 8419 (KANU). JEFFERSON 

CO.: P. H. Humfeld 201 (kanu). JOHNSON CO.: R. L. McGregor 
3431 (kanu). Leavenworth CO.: A. J. Petrik-Ott 750 (kanu). 
linn CO.: R. L. McGregor 34G5 (kanu). miamico.: R. L. McGregor 
2788 (kanu). Montgomery co. : L. C Hulbcrt 3678 (kanu). neosho 
co.: W. W. Holland 943 (kanu). osage CO.: R. L. McGregor 3316 
(kanu). shawnee CO.: A. J. Petrik-Ott 726 (kanu). wyandotte 
CO.: A. J. Petrik-Ott 748 (kanu). 



1975] Ferns — Petrik-Ott 489 



Nebraska: adams CO.: J. M. Bate*, 12 July 1913 (neb). CASS CO.: 
A. J. Petrik-Ott 111 (kanu). dotiglas CO.: H. J. Webber 6118 

(NEB). LANCASTER CO.: E. B. Robinson, May 189.", (neb). NEMAHA 

CO.: S. Steph ens 3626 (kanu). OTOE CO.: H. J. Webber 6115 (neb), 
richardson CO.: A. J. Petrik-Ott 775 (kanu). sarpy CO.: W. Cle- 
burne, 22 May 1888 (neb), tiiurston CO.: S. Stephens 21067 

(KANU). 

Cystopteris X tennesseensis Shaver, Journ. Tennessee 
Acad. Sci. 25(2) :107. 1950. 

Kansas: allen co.: S. Stephens 10688 (kanu). Anderson CO.: 
R L. McGregor 3457 (kanu). BOURBON CO.: A. J. Petrik-Ott 761 

(kanu). chase co.: F. Agrelht.8, 9 Aug. 1953 (kstc). Chautauqua 
co.: A. J. Petrik-Ott 744 (kanu). Cherokee co.: E. J. Palmer 
20*89 fKANU). clay co.: /. A very, 29 Sept. 1895 (ksc). coffey CO.: 
,/. E. Taylor, B. L. Taylor and L. K. Magrath 3820 (kanu). COWLEY 
CO • W. H. Horr 3152 (kanu). DOUGLAS Co.: A. J. Petrik-Ott 727 
(KANU). ELLSWORTH co.: R. L. McGregor 2696 (kanu). franklin 
CO.: R. L. McGregor 3095 (KANU). GREENWOOD CO.: R. L. McGregor 
17216 (KANU). Jefferson CO.: R. L. McGregor 3452 (kanu). 
johnson CO.: R. L. McGregor 3876 (kanu). Leavenworth CO.: 
S. Stephens 19444 (kanu). linn co.: A. J. Petrik-Ott 766 (kanu). 
Marshall CO.: S. Stephens 4665 (kanu). Miami CO.: P. H. Hmn- 
teld 212 (kanu). Montgomery CO.: A. J. Petrik-Ott 734 (kanu). 
NEOSHO CO.: S. Stephens 18763 (kanu). Osage CO.: 7?. L. McGregor 
3320 (kant'). Riley CO.: J. B. S. Norton, 25 Au ff . 1892 (mo). RUS- 
SELL CO.: W. H. Horr 3266 (kanu). SHAWNEE CO.: B. B. Smyth, 
May 1897 (KSC). wabaunsee co.: R. L. McGregor 12372 (kanu). 
wtlson CO.: R. L. McGregor 3404 (kanu). WOODSON CO.: ,4. J. 
Petrik-Ott 729 (kanu). WYANDOTTE CO.: 7?. L. McGregor 2418 

(KANU). 

Dryopteris carthusiana (Vill.) H. P. Fuchs, Bull. Soc 
France 105:339. 1958. 

Nebraska: brown CO.: F. Clements 2939 (neb). CASS CO.: E. 
Stoner, 14 May 1892 (neb), cherry co.: W. L. Tolstead 653 (neb). 

LANCASTER co! : H. J. Webber 6122 (neb). THOMAS CO.: P. A. 

Rydberg 1484 (neb). 

North Dakota: CASS co.: C. Wahlron, 5 July 1909 (nda). pem- 
bina co.: R. Brooks 457 (kanu). ransom Co.: R. A. Shunk, Aug. 
1916 (nda). richland co.: O. A. Stevens 1313 (nda). 

Dryopteris cristata (L.) A. Gray, Man. ed. 1. 631. 1848. 

Nebraska: hooker CO.: No collector given, 12 July 1893 (sdc). 
thomas CO.: P. A. Rydberg 1530 (neb). 



490 Rhodora [Vol. 77 



North Dakota: pembina CO.: R. Brooks 453 (kanu). ransom CO.: 
R. A. Shunk, Aug. 1916 (nda). 

Dryopteris filix-mas (L.) Schott, Gen. Fil. t. 9. 1834. 

South Dakota: CUSTER CO.: S. Stephens 5896 (kanu). Harding 
Co.: S. S. Visiter 7067 (sdu). Lawrence CO.: A. J. Petrik-Ott 794 
(kanu). meade co.: P. A. Rydberg 1197 (sdc). Pennington CO.: 
C. A. Taylor 8072 (sdc). 

Dryopteris marjrinalis (L.) A. Gray, Man. ed. 1. 632. 1848. 

Kansas: Anderson co.: E. Hartman 585 (kstc). Chautauqua 
CO.: A. J. Petrik-Ott 740 (kanu). Cherokee co.: R. L. McGregor 

984 (kanu). COFFEY CO.: R. L. McGregor 908 (kanu). elk CO.: 
R. L. McGregor 927 (kanu). greenwood co. : R. Brooks 469 and 
S. Stephens (kanu). Leavenworth co.: R. L. McGregor 2791 
(KANU). Montgomery CO.: R. L. McGregor 3394 (kanu). saline 
CO.: J. Hancin 1718 (ksc). wilson CO.: B. L. Wagenknecht 2252 
(kanu). woodson CO.: A. J. Petrik-Ott 733 (kanu). 

Gymnocarpium dryopteris (L.) Newm. Phvtologist 4: app. 
XXIV. 1851. 

South Dakota: custer co.: S. Stephens 5995 (kanu). Lawrence 
CO.: E. J. Pabner 37537 (umo). 

Phe^opteris hexagonoptera (Michx.) Fee, Gen. Fil. 243. 
1850-52. 

Kansas: Cherokee CO.: R. L. McGregor 3843 (kanu). 

Polystichum acrostichoides (Michx.) Schott, Gen. Fil. pi. 9. 
1834. 

Kansas: Chautauqua CO.: A. J. Petrik-Ott 741 (kanu). Cherokee 
CO.: S. Stephens 8589 (kanu). franklin CO.: S. Stephens 2890 



W 



MONT- 



GOMERY CO.: R. L. McGregor 949 (kanu). wilson co.: R. L. Mc- 
Gregor 922 (kanu). woodson CO.: R. L. McGregor 916 (kanu). 

Polystichum munitum (Kaulf.) Presl, Tent. Pterid. 83. 1836. 

South Dakota: Pennington co.: S. Stephens and R. Brooks 427 
(kanu). 

Thelypteris palustris Schott, Gen. Fil. t. 10. 1834. 

Kansas: Chautauqua Co.: R. L. McGregor 2414 (kanu). Doni- 
phan CO.: G. L. Clothier and H. N. Whitford, 31 July 1897 (ksc). 
Douglas CO.: R. L. McGregor 15065 (kanu). elk CO.: R. L. Mc- 
Gregor 928 (kanu). Ellsworth co.: O. A. Kolstad 1963 (kanu). 
greenwood cc: R. Brooks 468 and S. Stephens (kanu). mont- 



1975] Ferns — Petrik-Ott 491 



gomery CO.: R. L. McGregor 3397 (kanu). Pottawatomie co.: 

\V. A. Kellerman, 29 Sept. 1888 (ksc). riley co. : W. A. Keller-man, 

29 Sept. 1888 (mo), saline co.: J. Hancin 1761 (ksc). Washington 
CO.: T. C Dodd, Jr. 25 (ksc). wilson co. : R. L. McGregor 921 
(kanu). woodson CO.: R. L. McGregor 906 (kanu). 

Nebraska: BLAINE CO.: A'. Brooks 792 (kstc). brown CO.: C. E. 
Bissey, 23 July 1887 (neb), buffalo co. : Misses Smith and Lee 
7704 (neb), cherry co.: S. Stephen* 8201 (kanu). dodge CO.: 
C. E. Bessey, 9 Aug. 1872 (neb), douglas co. : W. Cleburne, Aug. 
18S1 (neb), franklin CO.: E. M. Hussong G777 (neb), hall CO.: 

R. J. Lemaire 2239 (neb), holt Co.: J. M. Bates, 31 July 1899 
(neb), jefferson CO.: A . F. Woods and D. Sounders 2004 (neb). 
Kearney CC: Dr. Hapeman, July 1892 (neb). LOUP CO.: S. Stephens 
6852 (kant). nemaha co.: ./. M. Bates, 5 July 1910 (neb). Sheri- 
dan CO.: F. Sandoz 378 (neb), thomas CO.: //. 7. Webber, 12 July 

1?89 (KSC). wheeler CO.: ft. L. McGregor 19370 (KANU). 

North Dakota: ransom CO.: R. A. Shunk, 15 Aug. 191 (> (nda). 

South Dakota: BENNETT co.: W. H. Over 15884 (sou). Penning- 
ton CO.: S. S. Visiter 1614 (sou), todd co. : T. Van Bruggen 5108 
(sdu). 

Woodsia obtusa (Spreng.) Torr. Geo. Rep. New York Nat. 
Hist. Sur. 4:195 Albany. 1840. 

Kansas: allen co.: P. H. Hum f eld 326 (kanu). Anderson CO.: 
P. H. Hum f eld 291 (kant;). atchison CO.: H. W. Blocker 809 (ksc). 
barber CO.: R. L. McGregor 14792 (kanu). barton CO.: S. Stephens 
8379 (kanu). bourbon CO.: A. J. Petrik-Ott 760 (kanu). CHAU- 
TAUQUA CO.: P. H. Humfeld 8(56 (kanu). Cherokee co.: A. J. 
Petrik-Ott 723 (kanu). clay CO.: C. Weber 318 (ksc). cloud CO.: 
S. V. Fraser 557 (ksc). COFFEY co. : G. L. Clothier and H. N. Whit- 
ford, 31 Aug*. 1897 (KSC). COWLEY CO.: Palmer 21256 (kanu). 
Crawford CO.: R. L. McGregor 976 (kanu). douglas Co.: R. L. 
McGregor 15074 (kanu). elk co.: S. Stephens 10767 (kanu). ells- 
worth CO.: R. L. McGregor 17325 (kanu). franklin co. : J. E. 
Bare 257 (kanu). greenwood co. : S. Stephens 15841 and R. Brooks 

(kanu). jefferson CO.: R. L. McGregor 883 (kanu). johnson 
CO.: R. L. McGregor 897 (kanu). labette co.: S. Stephens 11044 

(KANU). LEAVENWORTH CO.: A. J. Petrik-Ott 751 (kanu). LINCOLN 

CC: P. H. Humfeld 423 (kanu). linn CO.: A. J. Petrik-Ott 765 
(kanu). lyon co.: F. Agrelius, Sept. 1897 (kstc). Marshall Co.: 
S. Stephens 4664 (kanu). mcpherson co.: P. H. Humfeld 439 
(kanu). miami co.: P. H. Humfeld 222 (kanu). Montgomery co.: 

W. H. Horr E466 (kanu). nemaha CO.: 7?. L. McGregor 903 
(kanu). neosho CO.: S. Stephens 18763 (KANU). OSAGE CO.: R. L. 
McGregor 888 (kanu). Ottawa co.: S. Stephens 10913 (kanu). 



492 Rhodora [Vol. 77 



Pottawatomie CO.: P. H. Humfeld 393 (kanu). rice co.: E. L. 

Richards 1625 (kanu). RILEY CO.: P. H. Humfeld 406 (kanu). 
RUSSELL CO.: P. H. Humfeld 954 (kanu). saline CO.: P. H. Hum- 
feld 408 (kanu). shawnee co.: L. D. Voile 324A (kanu). Staf- 
ford CO.: A. H. Curtiss, 3 June 1905 (ksc). sumner CO.: D. Birk- 
holz 2785 (kanu). wabaunsee CO.: S. Stephens 4826 (kanu). 
Washington CO.: S. Stephens 10893 (kanu). WILSON CO.: R. L. 
McGregor 962 (kanu). woodson CO.: A. J. Petrik-Ott 719 (kanu). 
Wyandotte CO.: P. H. Humfeld 193 (kanu). 

Nebraska: Dakota CO.: 5. Stephens 21145 (kanu). douglas CO.: 
K. L. Johnson 1606 and D. E. Dallas (kanu). gage CO.: E. A. Fiala, 
7 Sept. 1925 (neb), hooker CO.: W. Kiener 24000 (mo), jefferson 
CO.: A. F. Woods and D. Saunders 2002 (neb). Lancaster CO.: 
A. S. Hunter 6774 (neb), pawnee CO.: C. H. Barnard, 1892 (neb). 
Saunders CO.: W. Cleburne, 26 July 1890 (neb), thurston CO.: 
S. Stephens 21119 (KANU). 

Woodsia oregana D.C. Eat. Can. Nat. and Geol. N.S. 2:89. 
1865. 

Nebraska: brown CO.: S. Stephens 24459 and R. Brooks (kanu). 
cherry CO.: W. L. Tolstead, 25 June 1937 (neb), custer CO.: A. J. 
Petrik-Ott 786 (kanu). DAwes CO.: S. Stephens 5671 (kanu). 
greeley CO.: R. Brooks 463 and S. Stephens (kanu). rock CO.: 
S. Stephens 6916 (kanu). scotts bluff CO.: W. L. Tolstead, 28 Au£. 
1941 (neb), sheridan CO.: S. Stephens 6188 (kanu). sioux Co.: 
/?. Brooks 475 and S. Stephens (kanu). Thomas CO.: P. A. Rydberg 
1479 (neb). 

North Dakota: billings CO.: R. Brooks 418 (kanu). golden 
valley CO.: S. Stephens 23439 and R. Brooks (kanu). mckenzie 
CO.: R. Brooks 416 (kanu). mclean CO.: J. Lunell, 18 June 1916 
(nda). morton CO.: R. Brooks 390 (kanu). sioux CO.: R. Brooks 
385 (kanu). 

South Dakota: CORSON CO.: S. Stephens and R. Brooks 383 (kanu). 
custer CO.: P. A. Rydberg 1199 (neb), fall river CO.: A. Nelson 
681 (sdu). Harding CO.: S. Stephens 7889 (kanu). lyman CO.: 
R. Brooks 376 and S. Stephens (kanu). Minnehaha CO.: L. J. 
Harms 2693 (kanu). Pennington CO.: R. L. McGregor 18786 
(kanu). Stanley CO.: R. Brooks 378 and S. Stephens (kanu). 
washabaugh CO.: J. E. Bare 1227 (kanu). 






Woodsia scopulina D.C. Eat. Can. Nat. and Geol. N.S. 2:91. 
1865. 

South Dakota: custer CO.: A. J. Petrik-Ott 785 (kanu). Law- 
rence CO.: E. J. Palmer 37556-A (umo). pennington co.: S. 
Stephens 7309 (kanu). 



1975] 



Ferns — Petrik-Ott 493 



enium 



Family Aspleniaceae 

platyneuron (L.) Oakes ex D.C. Eat. Ferns N. 
Amcr. 1 :24. 1878. 

Kansas: ATCHISON CO.: H. \V. Blocker (KSC). bourbon CO.: R.^L. 

McGregor 974 (kanu). CHAUTAUQUA CO.: A. J. Petrik-Ott 742 

(KANU). CHEROKEE CO.: A. J. Pelrik-OH 722 (KANT). COFFEY CO.: 

R. L. McGregor 909 (kanu). Crawford CO.: R. L. McGregor 988 

(KANU). DOUGLAS CO.: R. L. McGregor 660 (kanu). ELK CO.: R. L. 
McGregor 929 (kanu). franklin Co.: R. L. McGregor 899 (kanu). 

GREENWOOD CO.: R. L. McGregor 17208 (KANU). JOHNSON CO.: 
R. L. McGreoor 898 (KANU). LABETTE CO.: R. L. McGregor 9.. 8 

(KANU). LEAVENWORTH CO.: A. J. Petrik-Ott 749 (KANU). LINCOLN 

CO.: P. H. Humfeld 960 (kanu). MONTGOMERY CO.: A'. L. McGregor 
8396 (KANU). neosho CO.: R. L. McGregor 956 (kanu). WILSON 
ro. : R. J,. McGreqor 935 (kanu). woodson CO.: A. J. Petrik-Ott 

731 (KANU). 

Asplenium resiliens Kunze, Linnaea 18:331. 1844. 

Kansas: bourbon CO.: R. L. McGregor 986 (kanu). chautauqua 

CO ■ R. L. McGreqor 954 (kanu). CHEROKEE CO.: R. L. McGregor 
11090 (kanu). elk CO.: R. L. McGregor 938 (KANU). greenwood 
CO.: R. L. McGregor 936 (kanu). LABETTE CO.: R. L. McGregor 

969 (KANU). MONTGOMERY CO.: W. H. Horr E416 (kanu). WILSON 

CO.: R. L. McGregor 937 (kanu). 

Asplenium septentrionale (L.) Hoffm. Deutsch. Fl. 2:12. 
1795. 

South Dakota: custer CO.: S. Stephens and R. Brooks 429 (kanu). 
FENNINGTON CO.: R. Brooks All and S. Stephens (kanu). 

Asplenium trichomanes L. Sp. PL 2:1080. 1753. 

Kansas: CHAUTAUQUA CO.: A. J. Petrik-Ott 743 (kanu). green- 
wood CO.: R. L. McGregor 3416 (kanu). wilson CO.: F. Agrelius, 
28 May 1930 (ksc). WOODSON CO.: A. J. Petrik-Ott 730 (kanu). 

South Dakota: PENNINGTON CO.: P. A. Rydberg 1193 (neb). 

Asplenium viride Huds. Fl. Angl. 385. 1762. 

South Dakota: LAWRENCE CO.: S. Stephens and R. Brooks 426 
(kanu). 

Camptosorus rhizophyllus (L.) Link, Hort. Berol. 2:69. 
1833. 

Kansas: ALLEN CO.: R. L. McGregor 3336 (kanu). Anderson CO.: 
R. L. McGregor 3458 (kanu). atchison CO.: S. Stephens 10846 

(KANU). BOURBON CO.: A. J. Petrik-Ott 758 (KANU). CHAUTAUQUA 



494 Rhodora [Vol. 77 



co.: P. H. Hwmfeld 843 (kanu). Cherokee co.: R. L. McGregor 
11163 (kanu). coffey co.: R. L. McGregor 920 (kanu). cowley 
CO.: W. H. Horr and R. L. McGregor, 23 July 1947 (kanu). craw- 
ford co.: No collector given, no date (ksc). doniphan co.: F. 

Agrelius, Au ff . 1913 (kstc). DOUGLAS CO.: R. L. McGregor 29 
(kanu). elk CO.: R. L. McGregor 939 (kanu). franklin CO.: 
S. Stephens 17463 (kanu). greenwood Co.: W. H. Horr, 10 June 
1939 (kanu). Johnson co.: R. L. McGregor 2346 (kanu). la- 
bette co.: R. L. McGregor 959 (kanu). Leavenworth co.: W. M. 

Keith and B. L. Wagenknecht 2348 (kanu). linn CO.: A. J. Petrik- 

Ott 764 (kanu). miami co.: P. H. Humfeld 158 (kanu). Mont- 
gomery CO.: A. J. Petrik-Ott 737 (kanu). neosho CO.: R. L. Mc- 
Gregor 960 (kanu). riley co.: G. L. Clothier, 15 Sept. 1888 (ksc). 
wilson co.: R. L. McGregor 2370 (kanu). woodson co.: R. L. 
McGregor 919 (kanu). Wyandotte co.: M. Reed, 26 Sept. 1889 

(ksc). 



Family Blechnaceae 

Matteuccia struthiopteris (L.) Tod. Svn. PI. Acot. Vase 
Sicilia 30. 1866. 

North Dakota: bottineau co.: D. R. Moir, L. D. Potter and O. A. 
Stevens 2042 (nda). cass CO.: O. A. Stevens, 8 July 1922 (nda). 
Griggs CO.: No collector given, 11 June 1953 (nda). morton co.: 
O. A. Stevens, 14 Au«\ 1954 (nda). PEMBINA CO.: R. Brooks 450 
(KANU). ransom CO.: O. A. Stevens, 10 Sept. 1964 (nda). rich- 

lani, co.: W. J. Wanek 255 (nda). rolette co.: D. T. Disrud 98 
(nda). 

South Dakota: curter co.: S. Stephens 6105 (kanu). Penning- 
ton co.: S. Stephens 7117 (kanu). Roberts co.: A. J. Petrik-Ott 

799 (KANU). 

Onoclea sensibilis L. Sp. PI. 2:1062. 1753. 

Kansas: atchison CO.: H. W. Blocker 514 (ksc). CHAUTAUQUA 
CO.: R. L. McGregor 15046 (kanu). doniphan CO.: G. L. Clothier 
and H. N. Whiff ord, 31 July 1897 (ksc). douglas CO.: R. L. Mc- 
Gregor 704 (KANTJ). greenwood CO.: S. Stephens 9908 (kanu). 
Jefferson co.: R. L. McGregor 4152 (kanu). johnson CO.: J. A. 

Gummerwan, June 1888 (ksc). Leavenworth co.: P. H. Humfeld 

270 (kanu). Montgomery CO.: W. H. Horr and R. L. McGregor 
E421 (kanu). Ottawa CO.: C. T. Rogerson, 4 June 1954 (ksc). 
saline CO.: P. H. Humfeld 448 (kanu). woodson CO.: D. L. Marsh 

549 (KANU). 

Nehraska: brown CO.: S. Stephens 24433 and R. Brooks (kanu). 
buffalo CO.: Misses Smith and Lee, 1894 (neb), cherry co.: S. 



1975J Ferns — Petrik-Ott 495 



Stephens 8251 (KANU). franklin CO.: E. M. Huss&ng 6775 (neb), 
holt CO.: F. Clements 2802 (neb), jefferson CO.: R. A. Price 
7270 (neb), loup CO.: S. Stephens 6832 (kanu). otoeco.: C.Dunn, 

March 1904 (neb). SAUNDERS CO.: No collector given, 16 July 
1889 (rdc). sheridan CO.: F. Sandoz 366 (neb). THOMAS CO.: 
P. .4. Rydberg 1377 (neb). 

North Dakota: pembina CO.: O. A. Stevens 1594 (nda). RICHLAND 
CO.: O. A. Stevens 833 (nda). 

South Dakota: bennett co.: W. H. Over 15886 (sdu). custer 

CO.: S. Stephens 6016 (KANU). LAWRENCE CO.: A. C. Mcintosh 558 

(sou). Minnehaha CO.: W. H. Over 11102 (sdu). pennington co.: 
A. J. Petrik-Ott 793 (kanu). Stanley co.: H. E. Lee R154 (sdu). 



Family Dennstaedtiaceae 

Pteridium aquilinum (L.) Kuhn in v. d. Decken, Pveisen in 

Ost-Afrika3(3):ll. 1879. 

Kansas: Cherokee co. : R. L. McGregor 3863 (kanu). 

North Dakota: pembina CO.: R. Brooks 454 (KANU). 
South Dakota: Custer Co.: P. A. Rydberg 1192 (neb). LAWRENCE 
CO.: S. Stephens 7717 (kanu). 



Family Polypodiaceae 

Polypodium polypodioides (L.) Watt var. mithauxianum 

Weath., in Contr. Gray Herb. 124:31. 1939. 

Kansas: Chautauqua CO.: R. L. McGregor 3379 (kanu). 

Polypodium vulgare L. Sp. PI. 2:1085. 1753. 

South Dakota: custer co.: A. J. Petrik-Ott 790 (kanu). Minne- 
haha co.: L. ./. Harms 2688 (kanu). pennington co.: S. Stephens 
7227 (kanu). shannon CO.: S. S. Visiter, 6 July 1923 (sou). 



Family Marsileaceae 

Marsilea mucronata A. Br. Amer. Journ. Sci. 3:55. fig. 2. 
1847. 

Kansas: barber CO.: R. L. McGregor 10701 (kanu). barton co. : 
S. Stephens and R. Brooks 357 (kanu). butler CO.: P. H. Hum f eld 
798 (kanu). chare CO.: R. L. McGregor 3937 (kanu). clark co.: 
R. L. McGregor 4024 (kanu). clay CO.: S. Stephens 29475 (KANU). 
CLOUD CO.: S. V. Fraser 560 (ksc). COFFEY CO.: /. F. True, 22 July 

1938 (ksc). cowley CO.: S. Stephens 6583 (kanu). edwards co.: 



496 



Rhodora [Vol. 77 



W. A. Kcllerman, 21 Aug. 1884 (ksc). ELLIS CO.: R. L. McGregor 
3286 and W. H. Horr (kanu). finney co.: R. L. McGregor 3991 
(kanu). ford CO.: R. L. McGregor 3972 (kanu). GOVE CO.: A. S. 
Hitchcock, July 1895 (ksc). graham CO.: (7. L. Clothier, Au^.-Sept. 
1898 (KSC). gray CO.: R. L. McGregor 3270 and W. H. Horr 
(KANU). harper CO.: R. L. McGregor 13744 (KANU). HARVEY CO.: 
R. L. McGregor 3504 (kanu). HASKELL CO.: R. L. McGregor 4001 
(kanu). hodgeman CO.: R. L. McGregor 3962 (kanu). kingman 
co.: 7?. L. McGregor 7313 (kanu). kiowa CO.: /?. L. McGregor 
4951 (kanu). lane CO.: 7?. L. McGregor 3278 and TT T . 77. 7/V)?t 
(kanu). LYON co.: /. S. Wilson 9749 (kstc). meade co.: R. L. 

McGregor 3256 and TT 7 . 77. ffon- (kanu). morton co. : E. L. Ricltards 
3571 (kanu). osrorne Co.: J. Richardson and K. Robertson 856 

(KANU). PAWNEE CO.: /?. L. McGregor 3951 (kanu). POTTAWATOMIE 

co.: O. A. Stevens, 13 June 1909 (ksc). PRATT co.: TT 7 . 7\ Barker 
1809 (KANU). reno co.: E. W. Lathrop 2201 (kanu). republic co.: 

J. B. S. Norton, 23 Sept. 1895 (ksc). rice CO.: R. L. McGregor 3943 
(kanu). rush CO.: A. S. Hitchcock, Aujr. 1895 (ksc). saline co. : 
P. H Humfeld 871 (kanu). scott co. : A. S. Hitchcock, Aupr. 1895 
(ksc). sedgwick co.: R. L. McGregor 4047 (kanu). seward CO.: 
R. L. McGrenor 4010 (kanu). sheridan CO.: C. Weber 299 (KSC). 
sherman co.: ,4. S. Hitchcock, June 1892 (ksc). Stafford co. : 7. 
Jin ga r 629 (kanu). stanton co. : F. Lagergren. 20 Julv 1951 (ksc). 
STEVENS co.: R. L. McGregor 17015 (KANU). TREGO Co.: .7. L. 
Hutchison, 14 Aug. 1951 (ksc). Wallace co. : X. 5. Hitchcock, 
Aug*. 1895 (ksc). Washington CO.: R. L. McGrenor 1000 (kanu). 

wichita co.: Agrelius, Hall, Love joy 9 1913 (kanu). 

Nebraska: antelope CO.: A T . F. Petersen, June 1909 (neb), box 
butte co.: W. L. Tolstead, 28 Aug. 1941 (neb), chase co.: W. L. 
Tolstead, 4 Aug. 1941 (neb), clay co.: W. Kiener 22538 (neb). 

Custer CO.: /. M. Bates 2246 (neb), dawes co. : -7. M. Bates, 29 
July 1890 (neb), deuel CO.: E. M. Gilliard, 3 Julv 1893 (neb). 
DUNDY CO.: W. Kiener 10513 (neb), fillmore CO.: W. L. Tolstead, 

21 July 1941 (neb), franklin co.: W. L. Tolstead, 5 Sept. 1941 
(neb), gosper CO.: W. Kiener 19386 (neb), hall CO.: 7?. Brooks 

465 and S. Stephens (kanu). Hamilton co.: W. Kiefner 17847 
(neb), harlan CO.: W. L. Tolstead, 5 Sept. 1941 (neb), hayes 
co.: W. L. Tolstead, 7 Aug\ 1941 (neb). Jefferson co.: E. F. 

Large, 3 Sept. 1892 (neb). KEARNEY CO.: P. A. Rydberg 6604 
(NEE). Lancaster CO.: J. M. Bate*, 4 Au^. 1898 (neb), merrick 
CO.: I. Mueller, 14 July 1938 (neb), perkins CO.: W. L. Tolstead, 

7 Aup. 1941 (neb). PHELPS CO.: 7?. /. Lemaire 1499 (neb), pierce 

CO.: N. F. Petersen, 2 July 1907 (neb). SHERIDAN CO.: W. L. Tol- 
stead. 25 Aug:. 1941 (neb). WEBSTER CO.: <7. M. Bates, no date 
(neb), wheeler co.: 7?. Brooks 791 (kstc). 



1975] 



Ferns — Petrik-Ott 497 



North Dakota: barnes CO.: O. A. Stevens 1223 (NDA). BENSON 

co.: J. Lunell, 10 Sept. 1905 (nda). bottineau CO.: R. Smith, 20 

Sept. 1912 (NDA). bowman CO.: O. A. Stevens, 4 Julv 1949 (NDA). 
BURLEIGH CO.: S. Stephens 33268 and R. Brooks (KANU). foster 
CO.: 0. A. Stevens and W. A. Kluender, 21 Aup:. 1935 (nda). grant 
CO.: W. B. Bell 1355 (nda). redder CO.: 0. A. Stevens and D. R. 
Moir, 22 Aug-. 1956 (nda). MCLEAN CO.: O. A. Stevens, 27 Aug. 
1915 (NDA). MOUNTRAIL CO.: O. A. Stevens, 20 Aug. 1915 (nda). 
ramsey CO.: C. A. Geyer 71 (mo), slope CO.: A. C. Fox, 20 July 
1938 (nda). walsh CO.: A. C. Fox, July 1938 (neb), ward CO.: 

L. F. Lant.ev.sch lager 847 (nda). 

South Dakota: BROOKINGS CO.: T. A. Williams, Sept. 1893 (mo), 
brown CO.: A\ L. McGregor 18733 (kanu). BUTTE co.: A. Nelso'n 
549 (sou). CHARLES MIX CO.: J. Martin, 22 July 1905 (sou). CLARK 
CO.: Carter, Sept. 1896 (SDC). CUSTER CO.: A. Nelson, 2 Sept. 1920 
(sou), davison ro.: DiUman, 25 Aug. 1907 (sdc). HANSON CO.; 

7?. Brooks 435 (KANU). HARDING CO.: S. Stephens and R. Brooks 

423 (KANU). hughes co.: J. Martin, 24 July 1905 (sou), hyde co.: 

R. Brooks 400 (KANU). PERKINS CO.: S. S. Visher 7084 (SDU). SAN- 
BORN co.: W. H. Over, 3 June 1921 (sdu). Stanley co.: W. H. 

Over 7082 (sdu). WASHABAUGH CO.: S. S. Visher 7083 (sdu). 

Marsilea quadrifolia L. Sp. PI. 2:1099. 1753. 

Kansas: Cherokee CO.: O. A Kolstad and L. J. Harms 1581 
(KANU). neosho CO.: W. TT\ Holland 1993 (kanu). 

Pilularia americana A. Br. Monatsber. Akad. Berl. 1863: 
435. 1864. 

Kansas: HARVEY co. : R. Brooks 407 and S. Stephens (KANU). 



RENO CO.: R. L. McGregor 3933 (kanu). 

Nebraska: cherry CO.: R. L. McGregor 20017 (kanu). 



Family Azollaceae 

Azolla mexicana Presl, Abh. Bohm. Ges. Wiss. V. 3:150. 

1845 

Kansas: barber CO.: R. L. McGregor 14705 (kanu). barton co. : 
R. L. McGregor 5198 (kanu). Coffey CO.: L. K. Magrath 3291-1 

(kstc). douglas co.: R. L. McGregor 4687 (kanu). jefferson CO.: 

R. L. McGregor 4166 (kanu). linn CO.: O. A. Kolstad and L. J. 
Harms 2690 (kanu). LYON CO.: E. E. Garner 1110 (kstc). NEOSHO 
CO.: W. W. Holland 732 (kanu). 

Nebraska: FILLMORE CO.: W. Kiener 22552 (neb), garden CO.: 

W.Kiener2Sm (neb), keith co.: W. Kiener 23066 (neb), seward 

CO.: W. L. Totstead, 18 Aur. 1941 (neb). THOMAS CO.: A. J. Petrik- 
Ott 781 (kanu). 



498 Rhodora [Vol. 77 



THE FERN ALLIES 



Family Equisetaceae 

Equisetum arvense L. Sp. PI. 2:1061. 1753. 

Kansas: atchison CO.: R. L. McGregor 2807 (kanu). brown co.: 
Garner, Summer 1925 (ksc). clay CO.: Weber 409 (ksc). cloud 
CO.: S. V. Fraser 561 (ksc). doniphan CO.: A. J. Petrik-Ott 754 
(kanu). douglas Co.: R. L. McGregor 4234 (kanu). jackson 
CO.: R. L. McGregor 2844 (kanu). Jefferson CO.: R. L. McGregor 
4153 (kanu). johnson CO.: R. L. McGregor 3430 (kanu). leaven- 
worth co.: R. L. McGregor 3074 (kanu). Marshall co.: Stephens 
4701 (kanu). Pottawatomie co. : P. H. Humf eld 969 (kanu). riley 
CO.: F. C. Gates 15756 (ksc). saline Co.: J. Hancin 1255 (ksc). 

shawnee co.: B. B. Smyth 77 (ksc). wabaunsee co.: T. M. 

Barkley, 1 May 1963 (kanu). Washington co.: 7. C. Dodd, Jr. 
145 (ksc). woodson co.: E. W. Lathrop 2190 (kanu). wyandotte 
CO.: P. H. Humf eld 1021 (kanu). 

Nebraska: blaine co.: S. Stephens 24398 and R. Brooks (kanu). 
box butte Co.: C. H. Churchill, 5 July 1906 (neb), boyd co.: W. T. 
Barker 3059 (kanu). brown CO.: J. M. Bates, 25 June 1892 (neb), 
buffalo CO.: S. Stephens 6767 (kanu). burt CO.: S. Stephen* 32067 
and R. Brooks (kanu). cass CO.: S. Stephens 3768 (kanu). cedar 
CO.: R. Brooks 433 (kanu). cherry CO.: R. L. McGregor 19734 
(kanu). cuming co.: S. Stephens 36390 (kanu). dawes CO.: S. 
Stephens 5578 (kanu). dodge CO.: L. Magrath 2615 (kstc). DOUGLAS 
CO.: W. Cleburne, 23 May 1875 (neb), greely CO.: R. L. McGregor 
and J. E. Bare 430 (kanu). hall co.: R. J. Lcmaire 2064 (neb), 
holt co.: J. M. Bates, 22 June 1898 (neb), hooker CO.: S. Stephens 
17232 and R. Brooks (kanu). Howard CO.: J. E. Bare 390 and 
R. L. McGregor (kanu). keya paha CO.: S. Stephens 6934 (kanu). 
Lancaster Co.: Povml, Clements, and Saunders, 2 May 1893 (neb). 
LINCOLN CO.: S. Stephens 15995 and R. Brooks (kanu). LOGAN 
CO.: S. Stephens 25068 and R. Brooks (kanu). nance CO.: 
R. J. hemaire 1971 (neb), nemaha CO.: A. E. Watts, 5 May 1894 
(sou), otoe co.: Pound and Clements 4067 (neb). Richardson CO.: 
S. Stephens 3563 (kanu). sarpy CO.: Pound and Saunders 4097 
(neb), saunders CO.: J. E. Weaver, May 1936 (neb), seward CO.: 

H. J. Webber 6138 (neb). Sheridan CO.: S. Stephen* 6290 (kanu). 
Sherman CO.: R. L. McGregor 19801 (kanu). sioux CO.: S.Stephens 
16364 and R. Brooks (kanu). Thomas CO.: P. A. Rydberg 1378 
(neb), thurston Co.: S. Stephens 21118 (kanu). webster co.: 
7?. L. McGregor and J. E. Bare 792 (kanu). 

North Dakota: barnes CO.: H. F. Bergman 373 (nda). benson 
Co.: R. Brooks 457 (kanu). billings CO.: O. A. Stevens, 21 June 



1975] Ferns — Petrik-Ott 199 



1961 (NDA). BOTTINEAU CO.: R. Brooks 402 (KANU). BURKE CO.: 

R. Brooks 405 (kanu). cass Co.: H. F. Bergman, 17 June 1909 
(NDA). cavalier CO.: O. A. Stevens, 13 June 1956 (NDA). divide 
CO.: R. Brooks 407 (kanu). PUNN CO.: R. Brooks 397 (kanu). 
eddy CO.: 0. A. Stevens and I). R. Moir, 10 Sept. 1956 (nda). 
grand forks CO.: V. Facey, 14 July I960 (nda). GRIGGS Co.: 0. A, 
Stevens, 30 July 1965 (nda). LAMOURE CO.: J. F. Brenckle, 1905 
(nda). mchenry ro.: O. A. Stevens, 16 June 1956 (nda). mckenzie 
Co.: W. B. Bell 1038 (nda). mountrail co. : O. .4. Stevens and 
D. R. Moir, 13 Sept. 1956 (nda). Morton CO. : O. A. Stevens, 25 July 
1967 (nda). pembina CO.: R. Brooks 446 (kanu). ransom co.: 

W. B. Bell 290 (NDA). RICHLAND CO.: R. Brooks 445 (KANU). RO- 
LETTE CO.: D. T. Disrud 1952 (nda). SIOUX co. : A. J. Petrik-Ott 
796 (kanu). stark co.: O. A. Stevens, 17 Aug-. 1954 (nda). Steele 
CO.: O. A. Stevens, 7 July 1966 (nda). ward CO.: L. F. Lauten- 
schlager 425 (nda). Williams co. : R. Brooks 413 (kanu). 

South Dakota: Brookings co. : I>. Saunders, May 1897 (SDC). 
buffalo CO.: S. Stephens 32969 and R. Brooks (kanu). clay co. : 
W. H. Over 11012 (SDU). CODINGTON co.: ,/. R. Dugle 875 (sdu). 
Corson CO.: S. Stephens and R. Brooks 381 (kanu). CUSTER CO.: 
S. Stephens 6107 (kanu). deuel Co.: D. Saunders 8016 (sdu). 
hand co. : J. F. Todd 16820 (sdu). HARDING CO.: S. Stephens and 

7?. Brooks 424 (kanu). Lawrence Co.: S. Stephens 7578 (kanu). 
Lincoln CO.: L. ./. Harms 2604 (kanu). Marshall co. : T. Van 
Brvggen 446 (sdu). MINNEHAHA CO.: C. A. Taylor 9348 (SDU). 
moody co.: C. A. Taylor and R. Lowell 10251 (sdc). PENNINGTON 
co.: S. Stephens 7216 (kanu). Roberts co. : A. J. Petrik-Ott 798 
(kanu). sanborn co.: S. S. Visher 4145 (sdu). union co.: L. ./. 
Harms 2384 (kanu). YANKTON co.: L. J. Harms 2832 (kanu). 

Eouisetiim X ferrissii Clute, Fern Bull. 12:22. 1904. 

Kansas: Atchison co. : R. L. McGregor 2809 (kanu). barton 
CO.: R. L. McGregor 3944 (kanu). brown co. : IT. H. Horr, H) July 

1947 (kanu). Chautauqua co.: B. L. Waaenkneeht 1830 (kanu). 

Cherokee CO.: F. L. Richards 3124 (kanu). CHEYENNE CO.: R. L. 
McGregor 13582 (KANU). COWLEY co.: W. T. Barker 3657 (kanu). 
Doniphan CO.: B. L. Waaenkneeht 3341 (kanu). DOUGLAS CO.: 
F. B. Smith 412 (kanu). EDWARDS Co.: A. 5. Hitchcock, 1 Sent. 
1897 (ksc). Ellsworth co.: R. L. McGregor 12393 (kanu). jack- 
son co.: R. L. McGregor 2839 (kanu). jefferson CO.: Hartman 

1082 (KANU). Leavenworth co.: Hartman 1084 (kanu). LINCOLN 
CO.: P. H. Hum f eld 957 (kanu). linn co. : R. L. McGregor 3467 
(KANU). meade CO.: R. L. McGregor 4016 (kanu). Neosho CO.: 

IT". W. Holland 2506 (kanu). pottawatomie CO.: S. Stephens 4767 
(KANU). pratt co.: W. T. Barker 1574 (kanu). reno Co.: R. L. 



500 Rhodora [Vol. 77 



McGregor 12476 (kanu). republic CO.: D. K. Thomas, 1890 (ksc). 
riley CO.: No collector given, 1 July 1889 (mo), rooks CO.: E. 
Bartholomew, 25 June 1890 (neb), saline CO.: P. H. Humfeld 44fi 
(kanu). shawnee CO.: Hart man 1088 (kanu). trego CO.: M. Reed 
and A. S. <7. ? 8 July 1892 (ksc). wabaunsee CO.: S. L. Hunt 13 
(KSTC), Washington co.: W. H. Horr 4640 (kanu). Wichita co: 
F. Agrelius, no date £>iven (kstc). wilson co. : W. H. Haller, 
16 June 1896 (KSC). WOODSON CO.: S. Stephens 8383 (kanu). 
Wyandotte CO.: Hart man 1090 (kanu). 

Nebraska: brown CO.: S. Stephens 24439 and R. Brooks (kanu). 
buffalo CO.: R. L. McGregor and J. E. Bare 1656 (kanu). CASS CO.: 
S. Stephens 3771 (kanu). cedar CO.: S. Stephens 31978 and R. 
Brooks (kanu). cherry co.: S. Stephens 8167 (kanu). CUSTER 
CO.: J. M. Bates 2354 (neb). Dakota co. : S. Stephens 32054 and 
7?. Brooks (kanu). dawes Co.: H. J. Webber 6134 (neb), dixon 
CO.: F. Clements 2543 (neb), dodge CO.: L. Magrath 2616 (kstc). 
Douglas Co.: W. Cleburne, 8 July 1890 (neb), dundy CO.: A. F. 
Woods and D. Saunders 2000 (neb), hall CO.: 7?. J. Lemaire 2456 
(neb), holt CO.: J. M. Bates, 2 Aug. 1892 (neb). Kearney co.: 
P. A. Rydberg 475 (neb). SARPY CO.: W. Cleburne, 10 July 1897 

(neb), sioux co.: W. L. Tolstead, 13 Julv 19^0 (neb), stanton 

CO.: R. Brooks 432 (kanu). THOMAS CO.: H.'J. Webber 6131 (neb), 
valley CO.: S. Stephens 15620 and R. Brooks (kanu). w t ebster CO.: 
J. M. Bates, July 1903 (neb). 

North Dakota: barnes co. : //. F. Bergman 248 (nda). benson 
co.: J. Luncll, 15 Aug. 1915 (nda). cass co.: 0. A. Stevens 1083 
(nda). emmons CO.: R. Brooks 458 (KANU). GRIGGS CO.: 0. A. 
Stevens, 30 July 1965 (nda). MCHENRY CO.: O. A. Stevens, 21 July 
1966 (nda). MCINTOSH CO.: R. Brooks 459 (kanu). MORTON CO.: 
0. A Stevens, 6 Aug. 1938 (nda). pembina CO.: R. Brooks 451 
(kanu). ransom co.: W. B. Bell, 3 Julv 1909 (nda). richland 
co.: R. Brooks 444 (kanu). ward CO. : L. F. Lautenschlager 658 
(nda). Williams co.: 0. A. Stevens, 14 Aug. 1941 (nda). 

South Dakota: jackson co.: W. H. Over 7103 fSDU). meade co.: 
Snyder 307 (sdc). pennington co. : A. Nelson 514 (sdu). turner 
CO.: L. Messerli 532 (sdu). union CO.: L. J. Harms 2390 (kanu). 
WASHABAUGH CO.: S. S. Visher 207099 (sdu). 

Enuisetum fluviatile L. Sn. PI. 2:1062. 1753. 

Nebraska: Douglas CO.: E. R. Walker, 25 May 1940 (neb), 
garfield Co.: J. M. Bates 4917 (neb), holt CO.: J. M. Bates, 
27 June 1892 (neb), sarpy CO.: W. L. Tolstead 41594 (neb). 

North Dakota: benson co. : /. Lunell, 30 June 1908 (nda). 
bottineau co.: & Stephens 2884fi (kanu). mchenry CO.: J. Lu- 
nell, 24 June 1908 (nda). pembina CO.: L. R. Waldron 1604 (nda). 
richland CO.: 0. A. Stevens, 11 Julv 1934 (nda). rolette CO.: 



1975] Ferns — Petrik-Ott 501 



D. T. Disrud and D. Disrud 347 (nda). ward CO.: L. F. Lauten- 

sehlager 783 (nda). 

South Dakota: grant CO.: D. Saunders and J. P. Tawne, Aug. 
1897 (sdc). 

Equisetum hyemale L. var. afflne (Engelm.) A. A. Eat. 
Fern Bull. 11:111. 1903. 

Kansas: allen CO.: B. Osbom 64311 (ksc). ATCHISON CO.: P. L. 
McGregor 2733 (kanu). CHEROKEE CO.: R. L. McGregor 2734 

(kanu). clay CO.: C. Weber 316 (ksc). doniphan CO.: S.Stephens 
9349 (kanu). douglas co.: P. H. Humfeld 153 (kanu). Ellsworth 
co.: S. Stephens 17033 (kanu). franklin CO.: E. Hartman 644 
(KANU). GREENWOOD CO.: P. L. McGregor 12339 (kanu). JACKSON 
CO.: R. L. McGregor 2843 (kanu). JEFFERSON CO.: P. H. Humfeld 

138 (KANU). JOHNSON CO.: R. L. McGregor 3429 (kanu). LEAVEN- 
WORTH CO.: P. H. Humfeld 196 (kanu). linn CO.: S. Stephen* 
30999 (kanu). marion CO.: M. H. Burt el, 14 Oct. 1960 (KSC). 
Marshall CO.: S. Stephens 3316 (kanu). republic CO.: G. E. 
Morley 742 (kanu). riley CO.: F. C. Guirs and M. Newcomb, 
24 May 1928 (kanu). SUMNER CO.: G. L. Clothier and H. N. Whit- 
ford, 23 Au£. 1897 (ksc). Washington CO.: W. H. Horr 4640 
(kanu). wilson co.: H. Willis 7 (kstc). woodson co.: E. W. 
Lathrop 407 (kanu). wyandotte CO.: A. J. Petrik-Ott 746 (kanu). 
Nebraska: buffalo CO.: J. 7. Thornhrer 54 (neb), hurt CO.: 
S. Stevens 21036 (kanu). Cass CO.: J. L. Morrison 935 (neb). 

cherry CO.: IF. L. Tolstead 24 Aug. 1941 (neb), dawes CO.: W r . L. 
Tolstead, 25 Au#. 1941 (neb), douglas co.: W 7 . Cleburne, 8 Aug. 
1902 (neb), franklin co.: J. A7. j?afe.s, 9 June 1891 (neb), harlan 
CO.: P. L. McGregor 19531 (kanu). nemaha CO.: S. Stephens 17706 

(KANU). RICHARDSON CO.: A. J. Petrik-Ott 772 (KANU). SIOUX CO.: 

A. J Petrik-Ott 782 (kanu). THOMAS CO.: P. A. Rydberg 1722 
(neb), tiiurston co. : S. Stephens 21117 (kanu). 

North Dakota: CASS CO.: O. A. Stevens 2834 (nda). dickey CO.: 
7. F. Parr 860 and R. L. McGregor (kanu). eddy CO.: O. A. Stevens, 
13 Juno 1960 (nda). grand forks CO.: O. A. Stevens, 16 June 1953 

(nda). mchenry co.: O. A. Stevens, 16 June 1956 (nda). pembina 
CO.: O. A. Stevens and 7). P. /1/o/r, 13 June 1958 (nda). ransom 
CO.: S. Stephens 33487 and P. Brooks (KANU). RICHLAND CO.: 
A. P. Stoesz, 2 June 1934 (nda). 

South Dakota: CUSTER CO.: S. Stephens 5993 (kanu). DEUEL CO.: 
D. Saunders 8015 (SDC). HARDING CO.: S. S. Visher 7098 (SDU). 

Lawrence co.: S. Stephens 7424 (kanu). pennington co.: S. 
Stephen* 7217 (kanu). Roberts co.: P. Brooks 440 and 5. Stevens 
(kanu). shannon co.: S. S. Visher, 27 June 1916 (sdu). union 
CO.: P. E slick, 8 July 1961 (sdu). YANKTON CO.: S. Stephens 21229 

(kanu). 



502 Rhodora [Vol. 77 



Equisetum laevigatum A. Br. Amer. Journ. Sci. 46:87. 1844. 

Kansas: Anderson CO.: B. Neill 285 (kstc). barber co.: W. T. 
Barker 799 (kanu). barton CO.: H. A. Stephens, 12 June 1958 
(kstc). butler CO.: R. R. Weedon and L. K. Mag rath 4204 (kanu). 
Cheyenne CO.: H. A. Stephens, 14 June 1958 (kstc). clark CO.: 
R. L. McGregor 4025 (kanu). clay CO.: R. L. McGregor 2076 
(kanu). cowley CO.: D. E. Dallas 578 (kanu). decatur CO.: W. H. 
Horr 5046 (kanu). Dickinson CO.: Hartman 1064 (kanu). doni- 

pkan CO.: Hartman 1066 (kanu). DOUGLAS CO.: R. L. McGregor 
3027 (kanu). Ellsworth CO.: R. L. McGregor 17107 (kanu). 
finney CO.: R. L. McGregor 4000 (kanu). geary CO.: Hartman 

1072 (kanu). Hamilton CO.: E. L. Richards 3012 (kanu). harper 

CO.: E. J. Palmer 21201 (umo). Harvey CO.: W. H. Horr and R. L. 
McGregor 3724 (kanu). hodgeman CO.: R. L. McGregor 5188 
(kanu). jefferson co.: G. Latham 380 (kanu). kingman CO.: 

R. L. McGregor 7306 (kanu). kiowa CO.: R. L. McGregor and 
W. H. Horr 3832 (kanu). Leavenworth CO.: Hartman 1085 (kanu). 
logan CO.: S. Stephens 31297 and R. Brooks (kanu). marion CO.: 

P. H. Hum f eld 790 (kanu). Marshall co. : S. Stephens 3337 
(kanu). Mcpherson co.: L. /. Harms 1054 (kanu). meade CO.: 
W. H. Horr and R. L. McGregor 4053 (kanu). morton Co.: E. L. 
Richards 2499 (kanu). OTTAWA CO.: W. T. Barker 2545 (kanu). 
Pottawatomie CO.: Hartman 1086 (kanu). pratt co. : W. T. 
Barker 1611 (kanu). reno CO.: E. Hartman 889 (kanu). republic 
co.: G. E. M or ley 190 (kanu). rice co.: R. L. McGregor 12563 
(kanu). riley CO.: P. Zavos 29 (kstc). saline CO.: Hartman 
1074 (kanu). scott co.: R. L. McGregor 13007 (kanu). sedgwick 
CO.: M. A. Carleton, 27 May 1889 (ksc). shawnee CO.: Hartman 
1078 (kanu). Sherman CO.: O. A. Kolstad and L. J. Harms 1822 
(kanu). Stafford CO.: /. Vnga.r 511 (kanu). sumner CO.: J. W. 
Johnson 75 (kstc). trego CO.: S. Stephens 31211 and R. Brooks 
(kanu). Wallace CO.: R. L. McGregor 18579 (kanu). WICHITA 
CO.: Agrelius and Agrelius, 22 Aug 1 . 1912 (kanu). Woodson CO.: 
W. H. Horr, 10 July 1930 (kanu). 

Nebraska: boyd CO.: R. L. McGregor 19402 (kanu). box butte 
CO.: C. H. Churchill, 5 July 1906 (neb), brown co. : J. M. Bates, 
Juno 1893 (neb). BURT CO.: S. Stephens 32066 and R. Brooks 
(kanu). cass co.: S. Stephens 32146 and R. Brooks (kanu). cedar 
CO.: R. Brooks 434 (kanu). chase CO.: J. E. Bare 1403 and /?. L. 
McGregor (kanu). cherry co.: S. Stephens 8225 (kanu). custer 
CO.: A. J. Petrik-Ott 780 (kanu). Dakota CO.: S. Stephens 32035 
and R. Brooks (kanu). Dawes CO.: 5. Stephens 5565 (kanu). 
dawson CO.: R. L. McGregor and J. E. Bare 1619 (kanu). DIXON 
CO.: F. Clements 2542 (neb), dodge CO.: W. Kiener 29692 (neb), 
douglas CO.: W. Kiener 24571 (neb). DUNDY CO.: S. Stephens and 



197 5] Ferns — Petrik-Ott 50° 



o 



R. Brooks 358 (kanu). garden CO.: S. Stephens and R. Brooks 359 
(KANU). grant CO.: R. L. McGregor 10f582 (kanu). hall CO.: 

R. J. Lemaire 1559 (neb), hooker CO.: P. A. Rydberg 1801 (neb). 
Kearney co.: S. Stephens (5722 (kanu). KEinr CO.: S. Stephens 
and R. Brooks 361 (kanu). KEYA paiia CO.: S. Stephens 34319 and 

R. Brooks (kanu). knox CO.: S. Stephens and R. Brooks 373 

(KANU). LANCASTER CO.: W. Kiener 29597 (NEB). LINCOLN CO.: 

S. Stephens 15947 and R. Brooks (KANU). LOUP CO.: S. Stephens 
0901 (KANU). MADISON CO.: IT'. Kiener 29715 (mo). MCPHERSON 
CO.: S. Stephen? 2498(5 and R. Brooks (kanu). nance co.: B. 
Osborn 1108R (mo), nemaha co.: J. W. Gehling, 18 April 1935 
(SDU). otoe co.: B. Osborn 81 5R (mo), phelps CO.: S. Stephens 
42(58 and R. Brooks (KANU). RICHARDSON CO.: S. Stephens 3590 
(kanu). sarpy co.: W. Cb'bnrne, 21 Mav 1879 (ner). saunders 
CO.: T. B. Croat 2119 (kanu). scott bluff co.: S. Stephens and 
R. Brooks 3(53 (kanu). SHERIDAN co. : S. Stephens and R. Brook* 
430 (kant*). siiermax co.: TT\ T. Barker 2889 (kanu). SIOUX CO.: 
S. Stephens 1(5397 and R. Brooks (kanu). THOMAS CO.: R. L. Mc- 
Gregor 19651 (kant-). wkbster co.: R. L. McGregor 18042 (kant). 
wheeler co.: W. T. Barker 2955 (kant). 

North Dakota: barnes co.: O. A. Stevens 1218 (nda). BENSON 
CC: J. Lunell, 3 July 1910 (nda). billings Co.: R. Brooks 420 
(KANU). bottinfau CO.: O. A. Stevens, 15 July 19(50 (nda). bur- 
LEIGH CO.: .9. Stephens 33250 and R. Brooks (kanu). CASS CO.: Lee 
1205 (nda). dickey co.: J. E. Bare 859 and R. L. McGregor 

(KANU). DUNN CO.: R. Brooks 390 (KANU). EDDY CO.: O. A. 

Stevens, 13 July 1900 (nda). EMMONS CO.: O. A. Stevens 825 (nda). 

GOLDEN VALLEY CO.: O. A. Stevens, 11 July 1902 (NDA). GRAM) FORKS 

CO.: J. D. Walp, 19 June 1930 (nda). Gregory ro.: S. Stephens 
34071 and R. Brooks (kanu). hettinger co.: O. A. Stevens and 
TJ\ A. Kluender 137 (nda). lamoure CO.: ./. E. Bare 922 and R. L. 
McGregor (kanu). MCHENRY CO.: J. E. Bare 1039 and R. L. Mc- 
Gregor (KANU). MCKENZIE CO.: O. A. Stevcvs 1025 (NDA). MCLEAN 

CO.: V. Riid'!. 11 July 1958 (nda). mercer CO.: R. Brooks 393 
(kanu). MORTON CO.: S. Stephens 33014 and A'. Brooks (kanu). 
Mountrail CO.: J. E. Bare 1128 and R. L. McGregor (kanu). ran- 
som co.: O. A. Stevens, 7 July 1959 (nda). richland co.: O. A. 
Stevens 98 (nda). SARGENT CO.: H. L. Bulley, 10 June 1891 (nda). 
Sheridan CO.: O. A. Stevens, 9 July 1959 (nda). sioux CO.: R. 
Brooks 388 (kanu). slope CO.: R. Brooks 422 (kanu). stark CO.: 
O. A. Stevens, 17 Aug. 1954 (nda). STEELE co. : O. A. Stevens, 7 July 
1900 (nda). stutsman CO.: H. E. Bergman 89 (nda). ward CO.: 
R. Brooks 401 (kanu). Williams co.: O. A. Stevens, 20 June 1945 

(nda). 



504 Rhodora t Val - 77 



South Dakota: beadle CO.: C. A. Treadwell 104 (neb), brookings 
CO.: W. T. Barker 2697 (kanu). butte Co.: W. H. Over 17433 
(sdu). Campbell CO.: L. A. Hanna 128 (mo), clay CO.: W. H. 
Over 7104 (sdu). codington CO.: J. E. Dugle 109 (sdu). corson 
CO.: S. Stephens and R. Brooks 382 (kanu). Custer CO.: S. Stephens 
6064 (kanu). fall river CO.: S. Stephens 5790 (kanu). hamlin 
CO.: R. Brooks 436 and 8. Stephens (kanu). harding CO.: S. 

Stephens 7841 (kanu). Hutchison co.: V. L. Harms 454 (kanu). 
lawrence co.: S. Stephens 7401 (kanu). Lincoln co.: L. Mes- 
serli 821 (sdu). meade co.: S. Stephens 8097 (kanu). minnehaha 

co.: C*. A. 7«2/Zor 9349 (sdc). moody Co.: J. H. Wilde and C. A. 

Taylor 9287 (sdc). pennington co.: S. Stephens 7118 (kanu). 
perkins co.: S. Stephens 8029 (kanu). Roberts co.: R. Brooks 441 

and S. Stephens (kanu). SULLY CO.: S. Stephens 33077 and R. 
Brooks (KANU). TODD CO.: L. Stanley 199 (SDU). tripp CO.: S. 
Stephens 34119 and R. Brooks (kanu). union CO.: R. Eshck, 17 
Juno 1962 (sdu). washabaugh co.: S. S. Visher 7101 (sou), yank- 
ton CO.: L J. Harms 2533 (kanu). 

Equisetum palustre L. Sp. PI. 2:1061. 1753. 

North Dakota: ransom CO.: R. Brook* 1554 and S. Stephens 33495 

(kanu). 

Equisetum pratense Ehrh. Hannov. Mag. 22:138. 1784. 

North Dakota: pembina CO.: R. Brooks 448 (kanu). 

South Dakota: lawrence CO.: S. Stephens 7544 (kanu). pen- 
nington co.: R. Brooks 481 and 5?. Stephen* (kanu). Roberts co.: 

R. Brooks 438 (kanu). 

Equisetum scirpoides Michx. Fl. Bor. Amer. 2:281. 1803. 

South Dakota: LAWRENCE CO.: G. N. Jones and F. F. Jones 14862 
(MO). 

Equisetum sylvaticum L. Sp. PI. 2:1061. 1753. 

North Dakota: cavalier CO.: O. A. Stevens and D. R. Moir 1767 
(nda). PEMBINA CO.: R. Brooks 447 (kanu). 

South Dakota: Custer CO.: A. J. Petrik-Ott 791 (kanu). law- 
rence co.: E. J. Palmer 37347 (umo). pennington co.: C. A. 

Taylor 8112 (SDC). 



Family Isoetaceae 

Isoetes butleri Engelm. Bot. Gaz. 3:1. 1878. 

Kansas: CHEROKEE CO.: A. S. Hitchcock 1068 (ksc). 



1975] Ferns — Petrik-Ott 505 



Isoetes melanopoda Gay et Durieu, Bull. Soc. Bot. France 
11:102. 1864. 

Kansas: saline CO.: J. C. Hancin 2447 (kanu). Wilson CO.: 
R. L. McGregor 10267 (kanu). 

Nebraska: clay CO.: W. L. Tolstead, 5 July 1941 (neb), fill- 
more co.: W. L. Tolstead, 11 July 1941 (neb). Hamilton co.: 

W. Kiener 17851 (neb), harlan co.: W. Kiener 11850 (neb). 
KEARNEY CO.: W. L. Tolstead, 23 July 1941 (neb). 

South Dakota: MELLETTE CO.: W. H. Over 15878 (SDU). 



Family Lycopodiaceae 

Lycopodium obscurum L. Sp. PI. 2:1102. 1753. 

South Dakota: lawrence co.: P. A. Rydberg 1185 (neb) 



Family Selaginellaceae 

Selaginella engelmanii Hieron. Hedwigia 39(5) :294. 1900. 

Nebraska: holt CO.: W. T. Barker 2970 (kanu). 

North Dakota: billings CO.: R. Brooks 417 (kanu). bottineau 
CO.: O. A. Stevens, 25 July 1962 (nda). burke CO.: R. Brooks 406 
(kanu). dunn CO.: R. Brooks 394 (kanu). eddy CO.: O. A. Stevens 
2110 (nda). golden valley CO.: H. F. Berg-man 1158 (nda). 
grant Co.: Bell 632 (nda). mchenry co.: L. R. Haldron 2337 

(NDA). MCKENZIE CO.: R. Brooks 415 (KANU). MOUNTRAIL CO.: 
O. A. Steven* and D. R. Moir, 12 July 1960 (nda). siorx CO.: O. A. 
Stevens, 5 Auff. 1965 (nda). SLOrE CO.: R. Brooks 421 (kanu). 
stark co.: H. F. Bergman, 21 June 1910 (nda). ward CO.: J. Lunell, 
1 July 1909 (nda). WILLIAMS CO.: R. Brooks 412 (kanu). 

South Dakota: CUSTER CO.: P. A. Rydberg 1184 (neb). HARDING 

co.: Over and Solem 11358 (sdu). pennington Co.: S. Stephens 
7310 (kanu). perkins CO.: S. StepJien* 8106 (kanu). 

Selaginella rupestris (L.) Spring., Flora 21(12) :149, 182. 
1838. 

Kansas: CHAUTAUQUA CO.: A. J. Petrik-Ott 739 (kanu). douglas 
CO.: R. L. McGregor 4663 (kanu). elk CO.: R. L. McGregor 966 
(kanu). franklin CO.: R. L. McGregor 1003 (kanu). greenwood 
CO.: R. L. McGregor 3410 (kanu). Johnson CO.: R. L. McGregor 
1004 (kanu). Leavenworth CO.: R. L. McGregor 999 (kanu). 
Montgomery CO.: R. L. McGregor 996 (kanu). neosho CO.: R. L. 
McGregor 967 (kanu). wilson CO.: R. L. McGregor 995 (kanu). 
woodson CO.: A. J. Petrik-Ott 720 (kanu). 



506 Rhodora [Vol. 77 



Nebraska: antelope CO.: N. F. Petersen, 7 Aug. 1907 (neb), 
blaine CO.: S. Stephens 24376 and R. Brooks (kanu). box butte 
CO.: C. H. Churchill, about 1907 (neb), brown CO.: W. Kiener 29757 
(neb), cherry CO.: J. M. Bates, June 1891 (neb), holt CO.: J. M. 
Winter, 28 June 1931 (SDU). KEYA paha CO.: W. Kiener 23695 
(mo), loup CO.: S. Stephens 6858 (kanu). 

North Dakota: pembina CO.: R. Brooks 455 (kanu). 

South Dakota: codington CO.: W. H. Over 15474 (sou), custer 
CO.: A. J. Pctrik-Ott 788 (kanu). hanson co. : L. J. Harms 2759 

(KANU). MINNEHAHA CO.: L. J. Harms 2659 (KANU). PENNINGTON 

CO.: S. Stephens 7104 (kanu). turner co. : J. H. Martin 594 (sou). 



DOUBTFUL COLLECTIONS AND EXCLUDED TAXA 

Adiantum pedatum : This species was reported by Bessey 
(1892) for the Black Hills of South Dakota, but discounted 
by himself in his 1898 and 1900 publications. Rydberg 



(1917) 



pedatum 



in the Black Hills, although he states that the section in his 
manual concerning ferns was contributed by Miss Margaret 
Slosson. Miss Slosson, at the time of writing the fern sec- 
tion, was situated at the New York Botanical Garden. This 
leaves the possibility that there is a specimen, at the New 
York Botanical Garden, which I have not seen. 

Mcintosh (1949) reports the presence of Adiantum peda- 
tum in his work on the Black Hills, but his information is 
based upon Rydberg (1917), as is Winter, Winter, and 
Van Bruggen's (1959). Van Bruggen (1967), however, 
states that there is a specimen of Adiantum pedatum that 
was collected in Lawrence County, on the north slope of 
Spearfish Mountain by F. L. Bennett in 1941. Brooks 
(1969) states that this particular station has never been 
relocated. 

I have not seen the Bennett specimen or any other speci- 
mens of Adiantum pedatum from the Black Hills and, 
therefore, in my work, exclude it from this region. 

Asplenium trichomanes : There is a specimen of Asplen- 
ium trichomanes, located in the sdu herbarium and orig- 
inally misidentified as Cystopteris sp., from Devil's Lake, 
Ramsey Co., North Dakota, collected by W. H. Over, 22 



1975] Ferns — Petrik-Ott 507 



June 1908. The herbarium label accompanying this speci- 
men is quite confusing. The locality of collection written 
on the label is Devil's Lake, but the printed heading on the 
label reads, "Flora of South Dakota, State Geol. and Biol. 
Survey Collections, Plants of Clay Co., S.D.". There is no 
place in South Dakota known as Devil's Lake and therefore, 
someone later wrote in 'North Dakota' on the label. Devil's 
Lake is located in Ramsey County, North Dakota, but this 
record still remains questionable, due to the confusing label 
information and the lack of a suitable habitat in the area 
of Devil's Lake to support this species. 

Athyrium filix-f emina : Kansas: osage co. : C. H. 

Stokely, May 1895 (ksc). This specimen was collected in 
the town of Burlingame, Kansas. I would, however, be 
hesitant to accept this as a valid record since there is no 
suitable habitat near Burlingame where this fern might be 
found. Furthermore, it is commonly found in cultivation 
and C. H. Stokely is known to have collected many cultivars. 

Crypt oj? ram ma acrostichoides: Nebraska: franklin 
CO.: E. M. Hussong, 1893 (neb). The collection locality 
for this specimen is given as Franklin, Nebraska and no 
habitat information is given. It consists of only a few 
fertile segments. Cryptogramma acrostichoides is normally 
found in rocky crevices, on rocky slopes or slides. Such 
habitats are not to be found near the town of Franklin nor 
in the county. The closest collection site for this fern is in 
the Rocky Mountains of Colorado where it is common at 
altitudes of 7,000 to 12,000 feet. It would be difficult to 
accept this as a valid record for Franklin County, Nebraska. 

Dryopteris goldiana : Kansas : Leavenworth co. : J. 

Wilson, 1871 (kanu). This specimen was collected in the 
town of Leavenworth, Kansas. Mr. Wilson is known to 
have cultivated ferns and several of his cultivated speci- 
mens are on deposit in the KANU herbarium. This speci- 
men does not bear the abbreviation 'cult/ on the label 
although many of Wilson's other specimens do. McGregor 
and Hartman (1956) note: "It . . . seems as though the 



508 



Rhodora [Vol. 77 



labels were prepared by someone other than Wilson, for 
the same handwriting appears on other labels in the her- 
barium. It seems certain that the individual writing the 
labels merely omitted the 'cult/ inadvertently . . . , and 
thus introduced errors in our records." Wilson gives no 
habitat data or exact locality for his specimen. It is un- 
likely that there is any natural area near Leavenworth or 
anywhere in Kansas that would meet the habitat require- 
ments of Drvovteris aoldiana. 



urn 



reported many times as occurring in Kansas, Nebraska, 
South and North Dakota. I saw several specimens identi- 
fied to this species, but they were all depauperate forms of 
Equisetum lacvigatum. On the basis of the specimens I 
observed, this species is probably best excluded from the 
list of Equisetums for the four-state area. 

Matteuccia strut hiopteris: Cragin (1885) reports Mat- 
teuccia struthiopteris from Lawrence, in Douglas County, 
Kansas upon the word of Carruth. It is unlikely that it 
grows any place in Kansas in nature. I have seen it doing 
quite well in several gardens and greenhouses in Lawrence. 

Ophioglossum vulgatum: Gates (1940) reports Ophio- 
glossum vulgatum from Douglas County, Kansas upon the 
basis of a specimen at the University of Kansas. The kanu 
herbarium has no such specimen. Clausen (1938) cites a 
specimen collected in Crawford County, Kansas on June 
15, 1929 by F. A. Riedel and states that the specimen is on 
deposit in the herbarium of the New York Botanical Gar- 
den. Since I have not seen the specimen mentioned by 
Clausen (1938), or any duplicates, to check its validity, I 
am hesitant to include Ophioglossum vulgatum for Kansas 
until further evidence is secured. 

Osmunda claytoniana: This species has been erroneously 
reported from Nebraska by Rydberg (1932). 

Polystichum acrostichoides: Nebraska: No county 
given, L. Bruner, no date given (neb). No locality, other 



1975] Ferns — Petrik-Ott 509 



than Nebraska, or date of collection is given for this speci- 
men. It is probably best considered as a doubtful record. 

Thelypteris noveboracensis : Kansas: Pottawatomie 

CO.: G. Skimp, 30 April, no year given (KSC). This speci- 
men was collected near the town of St. George, Kansas. 
The habitat requirements of Thelypteris noveboracensis 
are not likely to be met in Kansas. Furthermore, Miss 
Shimp is known to have collected flowering plants from 
near St. George that are garden plants and its seems possi- 



pecimen 



similar habitat. 



Nebraska: douglas co. : W. Cleburne, 6 Oct. 1893 (neb). 

This specimen was collected in the city of Omaha, Nebraska. 
In the NEB herbarium is placed, also, a specimen collected 
on September 20, 1894, by W. Cleburne in a garden in 
Omaha. It is likely that the 1893 specimen was also col- 
lected in a garden and this information not mentioned on 
the label. 



ACKNOWLEDGEMENTS 



m 



cooperation of the curators at the following herbaria: 

KANU. KSC, KSTC, MO, NEB, NDA, SDC, SDU and UMO, who 

loaned me innumerable specimens for examination. The 
accumulation of the data presented here has also been 
greatly enhanced by financial assistance from the National 
Science Foundation Graduate Traineeship Program, The 
University of Kansas Committee on Systematics and Evo- 
lutionary Biology (GB-446X, Dr. George W. Byers, Prin- 
cipal Investigator) and the Sigma Xi Committee on Grants- 
in-Aid of Research. Special thanks are due my husband, 
Dr. Franklyn D. Ott, for his encouragement throughout the 
writing of this paper, Mr. and Mrs. Leo J. Petrik, Dr. 
Ronald L. McGregor, Mr. Ronald Weedon, Mr. Wayne 
Robuck, Mr. Kermit Johnson, and Miss Mary Wiegand who 
assisted me with field collecting, Mrs. Martha W. Rhodes 



manuscript, and M 



Ann Tyska 



manuscn 



510 Rhodora [Vol. 77 



LITERATURE CITED 



Bergman, H. F. 1918. Flora of North Dakota. Bienn. Rep. North 
Dakota Agric. Coll. Surv. 6: 151-372. 

Bessey, C. E. 1892. Ferns of the Black Hills. Amer. Nat. 26: 252- 
253. 



. 1898. The southern maidenhair fern in the Black Hills 

of South Dakota. Bot. Gaz. 26: 211. 

. 1900. One thousand miles for a fern. Asa Gray Bull. 



8: 2-6. 

Brooks, R. 1967. Ferns in Kansas. Kansas School Nat. 13: 1-15. 
■ •. 1969. The ferns of the Black Hills. Trans. Kansas 



Acad. Sci. 72: 109-136. 

Clausen, R. T. 1938. A ,mono^raph of the Ophio^lossaceae. Mem. 
Torrey Bot. Club 19: 5-171. 

Cragin, F. W. 1885. Notes on the ferns of Kansas. Bull. Wash- 
burn Lab. Nat. Hist. 1: 58-60. 
— - — . 1886. Further notes on Kansas ferns. Bull. Wash- 



burn Lab. Nat. Hist. 1: 175-176. 

Fernald, M. L. 1950. Gray's Manual of Botany. 8th Ed. American 
Book Co., New York, lxiv + 1632 pp. 

Fitzpatrick, T. J. 1919. The fern flora of Nebraska. Proc. Iowa 

Acad. Sci. 26: 311-326. 
. 1920a. The fern flora of Nebraska — I. Amer. Fein 



Jour. 10: 5-15. 



. 1920b. The fern flora of Nebraska — II. Amer. Fern 



Jour. 10: 33-44. 

Gates, F. C. 1940. Annotated List of the Plants of Kansas: Ferns 
and Flowering Plants (Flora of Kansas). Department of Botany, 
Kansas State College Contribution No. 391. 266 pp. 

HUMFELI), P. H. 1951a. A checklist of Kansas pteridophytes. Amer. 
Fern Jour. 41: 79-85. 

. 1951b. A checklist of Kansas pteridophytes. Amer. 



Fern Jour. 41: 79-85. 

KELLERMAN, W. A., & W. A. Kellerman. 1888. Analytical Flora 
of Kansas. Published by the Authors, Manhattan, Kansas. 197 
pp. 

Lanjouw, J. & F. A. STAFLEU (Eds.). 1964. Index herbariorum. 

Part I -—The Herbaria of the World. 5th Ed. Regyium Veg. 31. 
McGregor, R. L. 1960. Ferns and allies in Kansas. Amer. Fern 

Jour. 50: 62-66. 
, & E. L. Hartman. 1956. Notes of Kansas ferns. 



Amer. Fern Jour. 46: 84-87. 
MdNTOSH, A. C. 1949. A botanical survey of the Black Hills of 
South Dakota. Black Hills Eng\ 28: 3-74. 



1975} Ferns — Petrik-Ott 511 



Over, W. H. 1932. Flora of South Dakota. The University of 
South Dakota, Vermillion, S. D. 161 pp. 

Petersen, N. F. 1923. Flora of Nebraska. A List of the Ferns, 
Conifers, and Flowering Plants of the State with Keys for their 
Determination. 3rd Ed. Published by the Author, Plainview, 

Nebraska. 220 pp. 
Reed, M. 1 896. Ferns of Wyandotte County. Trans. Kansas Acad. 

Sci. 14: 150-51. 
Rydberg, P. A. 1917. Flora of the Rocky Mountains and Adjacent 
Plains. Published by the Author, New York, xii + 1110 pp. 

. 1932. Flora of the Prairies and Plains of Central 

North America. New York Botanical Garden, New York. 969 

pp. 
Saunders. D. A. 1899. Ferns and flowering plants of South Dakota. 

South Dakota Agric. Exp. Sta. Pull. No. 64. pp. 101-227. 
Stevens, O. A. 1950. Handbook of North Dakota Plants. North 

Dakota Institute for Regional Studies, Farpo, N. D. 324 pp. 
Van Bruggen, T. 1967. The pteridophytes of South Dakota. Proc. 

South Dakota Acad. Sci. 46: 126-144. 
Wilson, J. 1885. Annotated list of ferns found in Kansas. Pull. 

Washburn Lab. Nat. Hist. 1: r,6-r,8. 

cter, J. M., C. Winter, & T. Van Pritggen. 1959. A Checklist 

of the Vascular Plants of South Dakota. Department of Botany, 
State University of South Dakota, Vermillion, South Dakota. 
176 pp. 



Wi 



DEPARTMENT OF PLANT SCIENCES 
TEXAS A & M UNIVERSITY 
COLLEGE STATION, TEXAS 77843 



SEASONAL OCCURRENCE AND ECOLOGY 

OF MARINE ALGAE IN A 
NEW HAMPSHIRE TIDAL RAPID 1 



Norman B. Reynolds 2 and 
Arthur C. Mathieson 



Water motion is a major factor determining the growth 
and abundance of benthic plants in the ocean since it en- 
hances their metabolism (Conover, 1968) and assists in 
the propagation of plants and their dispersal to new areas. 
Several workers (see Lewis, 1964) have discussed the 
effects of wave action on the species composition and dis- 
tribution of seaweeds. Exposed coastal sites typically have 
more productive and diverse algal populations than shel- 
tered locations, presumably because of differential water 
motion. Sheltered embayments with strong tidal currents 
are also rich, productive habitats (Lewis, 1968; Schwenke, 
1971). To date, few detailed studies have been conducted 
of tidal rapid communities except for the comprehensive 
biological-hydrographical studies at Lough Ine, Ireland 
(Kitching and Ebling, 1967). In the present paper we 
describe the seasonal occurrence and ecology of seaweeds 
at the Dover Point, New Hampshire tidal rapid. 

The tidal rapid at Dover Point, New Hampshire, has 
several unique features which distinguish it from other 
tidal rapids. Foremost, it is located in the middle of the 
Great Bay Estuary System, rather than near the open 
coast (Fig. 1). Secondly, its substrate is stable even 
though it is partially composed of small pebbles and cob- 
bles. Thirdly, it has one of the fastest currents on the east 
coast of North America (Anon., 1969). 

Published with the approval of the New Hampshire Agricultural 
Experiment Station as Scientific Contribution Number 673. 

2 Present address: State University of New York, Cortland, N.Y. 



512 



1975] 



Marine Algae — Reynolds & Mathieson 



513 




Mass. 



42 55 



Fig. 1. The New Hampshire Coast and the Great Bay Estuary 
System. 



514 



Rhodora 



[Vol. 77 



MATERIALS AND METHODS 

Monthly collections of seaweeds were made during low 
tides from September, 1967, to September, 1969. Seasonal 
collections of sublittoral plants were made (by SCUBA) 
during- 1968. Diving- was only possible for about 20 min- 
utes at dead low tide; it was further restricted by the 
severe winter weather. No drift specimens are reported. 
All specimens were processed immediately after collection. 
Herbarium voucher specimens (a total of 1,264) were 
deposited in the Algal Herbarium of the University of 
New Hampshire (NHA). The specimens wer.3 identified 
according to Taylor (1957), with the exception of Fucus 
(Powell, 1957a, 1957b, 1963), Porphyra (Conway, 1964a, 
1964b), and Laminana (Wilce, 1965). The revised nomen- 
clature of Parke and Dixon (1968) was applied whenever 
possible. The longevity of the plants was designated ac- 
cording to Feldman's (1951) terminology. 

General weather conditions (rainfall, air temperatures 
and occurrence of ice) were recorded during field observa- 




Fig. 2. Dover Point and surrounding areas. 



1975] Marine Algae — Reynolds & Mathieson 515 



tions. Biweekly records of surface water temperatures, 
salinities and dissolved oxygen concentrations were moni- 
tored at dead low tide in the high and low current areas at 
Dover Point (Fig. 2). Salinities were recorded in the field 
with a set of hydrometers (G. M. Mfg. Co., New York) ; 
all of the values were corrected to 15°C. Temperatures 
were recorded with a common immersible thermometer. 
Oxygen concentrations were determined by use of a modi- 
fied Winkler method (Hach Chem. Co., Ames, Iowa). 
Diurnal variations of temperature, salinity, oxygen con- 
centration and current speed were recorded on five separate 
occasions. Salinities and temperatures were recorded at 
multiple depths with an Electrodeless Induction Sali- 
nometer (G. M. Mfg. Co., New York). The current speed 
was recorded with a Little Captain boat speedometer 
(Swift Instrument Company of Boston, Massachusetts), 
which was modified with a six-foot well tube. The accuracy 
of the instruments is ±: 1 /j, knots. 



DESCRIPTION OF AREA AND ENVIRONMENTAL FACTORS 

Dover Point is located at latitude 47°07'05" N. and longi- 
tude 70°49'50" W. in Dover, New Hampshire (Fig. 1). 
It is 5 miles northwest of Portsmouth, New Hampshire, 
and 5 miles east southeast of Durham, New Hampshire, at 
the junction of the Piscataaua River and Little Bay. All of 
the waters entering* and leaving- Little Bay, Great Bay and 
their five tributaries pass through the constricted channel 
at Dover Point, which is 470 yards wide and has a maxi- 
mum depth of 34 feet. 

The rock outcroppings at Dover Point are of the Eliot 
formation (Novotny, 1968). A variety of substrate types 
are present; they grade from boulders to cobbles, pebbles, 
sand and mud. The rocks are composed of mica schist with 
garnet crystals, phyllite, pegmatite-quarts, feldspar and 
metasiltstone. Smaller rocks and pebbles are stabilized by 
extensive mussel populations, which allow them to support 
relatively large plants. Most of the collections were ob- 



516 



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tained in the areas of high currents under the bridge 
(Fig. 2). The locations to the northeast and southwest of 
the bridge are mud flats which have reduced currents. The 
substrate in the high current area grades from rock near 
shore into mud at approximately 20 feet below mean low 
water. 

The tides at Dover Point are semi-diurnal, and they 
occur 87 minutes later than those of the adjacent open 
coast (Anon., 1965). The tidal amplitude is 6.8 feet. A 
maximum tidal current of 5.5 knots occurs 2.5 to 5 hours 
after low tide (Fig. 3). During ebb tide the current is 
concentrated on the Newington side of the channel; it is 
accompanied by a slight back eddy on the Dover Point side. 
Organisms in the lower intertidal and subtidal zones are 
exposed to strong currents at least 50% of the time (flood 
tide), while organisms in the upper intertidal zone are 
rarely exposed to strong currents. Subtidal organisms 
beyond the second piling (Fig. 2) are exposed to a nearly 
continuous current of high intensity. 

The annual range of water temperatures was —2.0° to 
19.0°C (Fig. 4). Diurnal temperature fluctuations at Dover 
Point depend on the two water masses involved — i.e., 
Great Bay and the Atlantic Ocean. Little or no diurnal 
variation (0-0. 5°C) occurred during the winter. The larg- 
est diurnal variation (5°C) occurred during the late spring 
and summer. The annual range of salinity at Dover Point 
was 7.5 o/oo at spring runoff to 31.0 o/oo in the late sum- 
mer (Fig. 4). It usually varied from 23-29 o/oo. The 
greatest diurnal fluctuation of salinity was recorded during 
March, 1969 (i.e., 10 o/oo). In general there was a decrease 
in salinity from low tide until one hour after the predicted 
low. Thereafter it rose until high tide at which time it 
decreased for two to three hours. The dissolved oxygen 
concentration varied seasonally, with peak values occurring 
in the spring (12-14 ppm) and minimal valu 



summer (6-8 ppm 



temper 



salinity and oxygen values were found between the high 
and low current areas. 



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1975] Marine Algae — Reynolds & Mathieson 519 



SPECIES COMPOSITION 

The algal flora at Dover Point shows a high diversity of 
species for an estuarine habitat. Eighty-three taxa of sea- 
weeds were recorded from the tidal rapid site at Dover 
Point, including 39 Rhodophyceae, 24 Chlorophyceae and 
20 Phaeophyceae. Table I summarizes the number and 
kind of seaweeds encountered at Dover Point as well as at 
adjacent coastal and estuarine locations. Seventy-one of 
the taxa found at Dover Point were also found on the ad- 
jacent open coast (Mathieson & Fralick, 1972; Maithieson, 
Hehre & Reynolds, in press). In contrast, only 59 taxa 
from Dover Point were found in adjacent estuarine sites 
with calm waters (Mathieson, Reynolds & Hehre, in press). 
The red and brown algae at Dover Point showed more af- 
finities to the open coast than the green algae (Table T). 

SEASONAL OCCURRENCE 

Tables II-IV summarize the monthly occurrence and 
longevity of each species based on three years of monthly 
collections at Dover Point. The m 

cies of Rhydophyceae were found in July (Table V). Peak 
numbers of Chlorophyceae were evident in May, while the 
Phaeophyceae showed peaks in April and May (Table V). 
Table V also records the total number of seaweeds collected 
per month; the largest number of species occurred during 
May to July. The low numbers recorded in January and 
September resulted from a lack of subtidal collections. 

Several taxa were rare at Dover Point. Brijopsis plu- 
mosa, Enteromorpha compressa, Monostroma leptodermum, 
Ascophyllmn nodosum forma scorpioides, Fucus distichus 
spp. cvanenscens, Myrionema strangulans, Sphacelaria cir- 
rosa, Ceramium strictum, Dermatolithon pustulatum, Glo- 
iosiphonia capillaris, Melobesia lejolisn and Rhodoehorton 
purpureum were only found once during the entire three 
year period. Of particular interest was the sporadic oc- 
currence of Gloiosiphonia capillaris, for it was only seen 
during a two week period and it was represented by 10 
plants within a 10 ft- area. A comparison of Tables II-IV 



iximum 



520 Rhodora [Vol. 77 



shows that six of the above species are annuals, which 
might be expected to have a sporadic distribution. Other 
seaweeds such as Callithamnion baileyi, Dasya pedicellata, 
Lomentaria orcadensis, Polysiphonia harveyi, P. lanosa 
and Laminaria longicruris were collected twice during the 
36 month study. Monostroma pulchrum, Gracilaria foli- 
ifcra, Porphyra miniata and Ckordaria flag elliformis were 
collected three times. Laminaria longicruris, Gracilaria 
foliifera and Polysiphonia lanosa are perennials, while all 
the others are annuals (Tables III and IV). 

Forty-two of the 83 taxa collected at Dover Point were 
considered to be perennials (Table VI). Most of the green 
algae were annuals (79%). A larger portion of the brown 
(65%) and red algae (62%) were perennials. Two dis- 
tinct types of annuals are present at Dover Point : seasonal 
and aseasonal. Seasonal annuals occur during a certain 
season and subsequently disappear. Monostroma grevillei, 
Spongomorpha arcta, Bangia fuscopurpurea, and Urospora 
penicilliformis are examples of late winter and/or spring 
annuals. Chorda tomentosa, Bryopsis plumosa, Antitham- 
nion cruciatum, Callithamnion baileyi, Ceramium strictum 
and Dasya pedicellata are summer annuals. Aseasonal an- 
nuals, such as Ectocarpus confervoides, Petal onto fascia, 
Scytospihon lomentaria, Enteromorpha intcstinalis and 
Ulothrix flacca occur all year long, although they may have 
peaks in the spring and summer. Aseasonal annuals are 
represented by several generations of plants. 

Two major types of perennial algae are also evident at 
Dover Point. Ascophyllum nodosum, Fucus vesiculosus, 
Laminaria spp., Pseudendoclonium marinum, Ahnfeltia 
plicata, Chondrus crispus, Gigartina stellata, Phyllophora 
membranifolia and Rhodymenia palmata are typical of the 
most common type where the whole plant is perennial. 
Other species such as Elachista fucicola, Pilaiella littoralis, 
Cladophora sericeae and Phycodrys rubens perenniate by 
a portion of the thallus. Transitional forms are also evi- 
dent between the two types. Knight and Parke (1931) 
designate plants as pseudperennials if a small persistent 
portion of the thallus can regenerate the intact plant. 



1975] Marine Algae — Reynolds & Mathieson 521 



DISCUSSION 

The algal flora at Dover Point is "open coastal" in char- 
acter for it has a very productive and diverse flora, as well 
as a large number of species in common with the open 
coast. It should be emphasized that adjacent, estuarine 
areas, that lack currents but have similar substrate and 
hydrographic conditions support a less diverse and reduced 
vegetation. Lewis (1964) reports a similar biological char- 
acterization of tidal rapids in Scotland and Ireland. In 
addition he records the "dying out" of more delicate algae 
in favor of more robust forms with increased water flow. 

Kitching and Ebling (1967) state that the major in- 
fluence of tidal currents is exerted on the sublittoral zone. 
They also suggest that tidal currents are analogous to wave 



lmer 



extremes 



mechanical pull. Moore (1966) states that tidal rapids 
support open coastal invertebrates, even though the loca- 
tions are essentially sheltered. The sublittoral zone at 
Dover Point has more "open coastal" species than the 
littoral zone. 

Pronounced seasonal fluctuations of algal species were 
recorded at Dover Point, with the largest number of spe- 
cies occurring in July and the lowest in the winter. The 
wide range of hydrographic conditions, particularly tem- 
perature, probably causes the seasonal differences in its 
flora. Williams (1948, 1949) and Coleman and Mathieson 
(1975) have also recorded a wide range of annuals in areas 
with pronounced temperature fluctuations. A combination 
of boreal (e.g., Dumontia incrassata, Porphym umbilicalis) 
and warm temperate annuals (e.g., Callithamnion bailey i, 
Dasya pedicellata) are present in the Great Bay Estuary 
System, because of the wide range of temperatures ( Math- 
ieson, Reynolds and Hehre, in press). The seasonal occur- 
rence of organisms at Dover Point is very similar to that 
at the adjacent open coast near Portsmouth, New Hamp- 
shire (Mathieson, Hehre & Reynolds, in press), except that 
a larger number of spring annuals is evident during April 
on the open coast. 



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Marine Algae — Reynolds & Mathieson 



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1975] Marine Algae — Reynolds & Mathieson 531 



TABLE v. 

NUMBERS OF TAXA OF RHODOPHYCEAE, FHAEOPHYCEAE, 
AND CHLOROPHYCEAE COLLECTED AT DOVER POINT 

DURING VARIOUS MONTHS, 1967-1969 

JFMAMJ JASOND 

Rhodophyceae 19 19 21 19 25 24 27 22 15 21 20 21 

Phaeophyceae 10 13 13 16 15 14 13 11 11 13 11 12 

Chlorophyceae 9 11 12 13 15 12 13 11 12 14 11 12 

Total 38* 43 46 48 55 50 53 44 38* 48 42 45 

*No subtidal collections were made 



TABLE VI. 

NUMBERS OF PERENNIAL RHODOPHYCEAE, 
PHAEOPHYCEAE AND CHLOROPHYCEAE AT DOVER POINT, 

1967-1969 

Number of taxa Total ( /r of total % of 

potentially Number taxa which perennial 
perennial of taxa were perennial taxa/class 

Rhodophyceae 24 39 29 62 

Phaeophyceae 13 20 16 65 

Chlorop hyceae 5(7) 24 6(8) 21(29) 

Grand Total 42(44) 83 61(64) 49(52) 



ACKNOWLEDGEMENTS 



We would like to thank Dr. A. Hodgdon for his critical 
review of the manuscript. In addition we express our grati- 
tude to Drs. William Flahive and Richard Burns for as- 
sistance in the collection of field data. 



LITERATURE CITED 



Anon. 1965. Tide tables, high and low water prediction, east 
coast of North and South America, including Greenland, 1966. 
U. S. Dept. of Commerce, Coast and Geodetic Survey. 289 pp. 

Washington, D.C. 
. 1969. Tidal current tables, Atlantic coast, North 



America, 1969. U. S. Dept. of Commerce, Coast and Geodetic 
Survey. 200 pp. Washington, D.C. 



532 



Rhodora [Vol. 77 



Coleman, D. C., & A. C. Mathieson. 1975. Investigations of New 
England marine algae VII: Seasonal occurrence and reproduc- 
tion of marine algae near Cape Cod, Massachusetts. Rhodora 
77: 76-104. 

Conover, J. T. 1968. The importance of natural diffusion gradients 
and transport of substances related to benthic marine plant 
metabolism. Bot. Mar. 40: 1-9. 

Conway, E. 1964a. Autecological studies of the genus Porphyra: 

I. The species found in Britain. Brit. Phycol. Bull. 2: 342-346. 
. 1964b. Autecological studies of the genus Porphyra: 

II. Porphyra umbilicalis (L.) Jour. Ag. Brit. Phycol. Bull. 2: 
349-363. 

Feldmann, J. 1951. Ecology of marine algae. Pp. 313-334 in: 
G. M. Smith (ed.) Manual of Phycology. Ronald Press, New 
York. 

Kitching, J. A., & F. J. Ebling. 1967. Ecological studies at Lough 

Ine. Adv. Ecol. 4: 197-291. 
Knight, M., & M. W. Parke. 1931. Manx algae. Mem. Liverpool 

Marine Biol. Comm. 30: 1-147. 
Lewis. J. R. 1964. The ecology of rocky shores, xii + 323 pp. 

English Univ. Press Ltd., London. 
. 1968. Water movements and their role in rocky shore 



ecology. Ecology. Sarsia 34: 13-36. 

Mathieson, A. C, & R. A. Fralick. 1972. Investigations of New 

England marine algae V. The algal vegetation of the Hampton- 
Seabrook Estuary and the adjacent open coast near Hampton, 
New Hampshire. Rhodora 74: 406-435. 

, E. Hehre, & N. B. Reynolds. Investigations of New 

England marine algae I. A floristic and descriptive ecological 
study of the marine algae of Jaffrey Point, New Hampshire. 

Nova Hedwigia, in press. 
, N. Reynolds, & E. Hehre. Ibid II. The species com- 



position, distribution and zonation of seaweeds in the Great 
Bay Estuary System and the adjacent open coast of New 
Hampshire. Nova Hedwigia, in press. 

Moore, H. B. 1966. Marine Ecology, xi + 493 pp. John Wiley & 
Sons, New York. 

NOVOTNY, R. N. 1968. Geologic Map of the Seacoast Region, New 
Hampshire Bedrock, N.H. Dept. Resources and Econ. Develop- 
ment. 

Parke, M., & P. S. Dixon. 1968. Checklist of British marine 
algae — second revision. Jour. Mar. Biol. Ass. U.K. 48: 783-832. 

Powell, H. T. 1957a. Studies in the genus Fucus L. I. Fueus 
distichus L. emend. Powell. Jour. Mar. Biol. Ass. U.K. 36: 407- 
432. 



1975] Marine Al^ae — Reynolds & Mathieson 533 



. 1957b. Studies in the genus Fucm L. II. Distribu- 
tion and ecology of forms of Fucks- distichus L. emend. Powell 
in Britain and Ireland. Jour. Mar. Biol. Ass. U.K. 36: 663-793. 
. 1963. Speciation in the genus Fucus L. and related 



genera. Pp. G3-77 in: J. P. Harding & N. Tebble, eds. Specia- 
tion in the sea. System. Ass. Publ. No. 5. 

Schwenke, H. 1971. Water movements II. Plants. Pi). 1091-1121 
in: 0. Kinne, ed. Marine Ecology. Vol. I, Pt. 2. 

Taylor, W. R. 1957. Marine Algae of the Northeast coast of 
North America, viii + 509 pp. Univ. of Michigan Press, Ann 
Arbor. 

Wilce, R. T. 1965. Studies in the genus Laminaria. III. A re- 
vision of the North Atlantic species of the Simplices Section of 
Laminaria. Botanica Gothberg. 3: 247-256. 

Williams, L. G. 1948. Seasonal alternation of marine floras at 
Cape Lookout, North Carolina. Am. Jour. Rot. 35: f>82-f>95. 

. 1949. Marine algal ecology at Cape Lookout, North 



Carolina. Bull. Furman Univ. 31: 1-21. 



DEPARTMENT OF BOTANY AND PLANT PATHOLOGY 
AND JACKSON ESTUARINE LABORATORY 
UNIVERSITY OF NEW HAMPSHIRE 
DURHAM, NEW HAMPSHIRE 03824 



BOOK REVIEW: 
THE FLORA OF ESSEX COUNTY, 

MASSACHUSETTS 1 



James P. Poole 



In 1880 a Flora of Essex County, Massachusetts was 
published by John Robinson who had been in charge of the 
herbarium of the Peabody Museum in Salem, Massachusetts 
for a number of years, eventually becoming Director of 
the Museum. His flora consisted of a list of the species 
then growing or reported to be growing in the County, 
often with the names of the collectors but with only scant 
information concerning the habit or habitat or the localities 
where the species were found. The genera were listed 
under families that were referred to as "orders" in the 
summary, following the fifth edition of Gray's Manual of 
Botany. That publication, in turn, followed DeCandolle as 
modified by Bentham and Hooker's Genera Plantarum. In 
addition to the vascular plants, listed as Exogens, the flora 
flso included mosses, liverworts, lichens, and algae; the 
fungi were covered by only a brief reference. 

In 1951 Stuart Harris, then Professor of Biology at Bos- 
ton University, acceded to a request to write a modern 
flora of the County for publication by the Peabody Museum. 
From that date on he devoted what time he could spare 
from his teaching duties and other varied interests to in- 
tensive study in the field and in some of the herbaria in 
which early collections from Essex County had been de- 
posited. After many years he completed the Catalog of the 
Flora and some of the accompanying sections, but unfortu- 
nately did not live to complete a preface or write some of 
the introductory sections he had planned. These portions 
have been supplied by others. Dorothy Eastman Snyder, 

^The Flora of Essex County, Massachusetts by Stuart Kimball 
Harris. Peabody Museum, Salem, Massachusetts. 1975. vii + 269 pp. 
Map of the County on the inside cover. Sketch of Sweet Bay flower 
on title page. $12.50 



534 



1975] Book Review 535 



Consultant in Natural History at Peabody Museum, with 
Stuart Harris, authored the preface and also served as 
Editor. Other sections included: a Foreword by Richard 
Jefferson Eaton, former Curator of the herbarium of the 
New England Botanical Club, who also contributed a brief 
biographical sketch of the author; one on Geology in Rela- 
tion to the Flora by Sarah Fraser Robbins, Director of 
Education at the Museum; another by Robert Lincoln 
Goodale, M.D., entitled History of Botanical Collecting, 
with biographical information about some of the collectors 
of Robinson's time and earlier; and a short section on 
Climate by Hurd C. Willett, Professor Emeritus of the 
Department of Meteorology, M.I.T. An End Map was 
drawn by Charles F. Allen IT, Assistant at the Museum. 
The sections contributed by Dr. Harris included: a Catalog 
of Species; a Statistical Summary of genera, species, vari- 
eties, forms, and hybrids listed by families with each group 
of taxa designated as Native or Introduced; a list of Spe- 
cies Exclusae; and a list of collectors named in the Cata- 
log with information as to the period and area of activity 
of each of them. A list of references is also included. 

Tn the Catalog the arrangement of the families and the 
nomenclature follow Gray's Manual, 8th edition, 1950, with 
few exceptions, but the specific epithets are in lower case 
as recommended in the International Code of Botanical 
Nomenclature, 1972. Under the genera the species are 
listed with both Latin and common names, with a brief 
statement as to habitat and frequency, followed by the 
names of towns or cities where collected, the names of the 
collectors, with collectors' numbers and date of collection 
when available. For the varieties, forms and hybrids, only 
the locality where collected, the collectors, collectors' num- 
bers, and dates are given, except that information as to 
habitat and frequency is included for hybrid species. Pre- 
ceding the Catalog is a list of towns and cities numbered 
in the order quoted in the Catalog. These numbers are 
spotted on the End Map. 



536 Rhodora [Vol. 77 



In the text there are a few misstatements and typo- 
graphic errors, and in the Catalog there are a number of 
errors, mostly typographic. In the list of references and 
also in the section listing the publications by the author 
there are numerous mistakes, mostly in the year or the 
volume number of the publication cited. In addition there 
are some discrepancies which should be recorded. A few 
of the species included in the Species Exclusae are also 
listed in the Catalog. It seems probable that Dr. Harris 
may have decided that some of the questionable species 
should be included in the Catalog but failed to find the time 
later to delete the names from the Species Exclusae. Again, 
if the Statistical Summary is checked against the Catalog, 
it will be discovered that there are instances in which the 
two do not agree. Any corrections in the Statistical Sum- 
mary must affect the Total Summary by Minor Groups on 
page 23. Thus it is impossible to make any exact compari- 
son in totals with Robinson's Flora. In Harris's Flora the 
figure given for the Grand Total Taxa (1767) is the same 
number as that given for the total species. It is evident 
that it fails to include the totals for varieties, forms, and 
hybrids. 

These discrepancies and errors are difficult to explain 
but, knowing the competence of Stuart Harris and his 
scholarly standards, it seems most probable that the incon- 
sistencies and most of the other errors would have been 
remedied if he could have lived to make that final check 
that is so essential before any manuscript goes to the 
printer. 

Fortunately, none of these difficulties or errors are of 
enough significance to detract from the interest or value of 
this publication. The catalog of species constitutes an 
authoritative checklist of the flora of the County and, as 
emphasized in the Foreword (p. vii), Essex County is note- 
worthy from a phytogeographical point of view. It is an 
area where Canadian zone species overlap traces of a 
southern flora, where the northern Habenaria blephari- 
glottis grows in the shelter of an indigenous colony of 



1975] Book Review 537 



Magnolia virginiarm, Moosewood {Acer pensylvanicum) 
occurs rather abundantly within a few rods of the sea-cliffs 
at Manchester, and Hobble-bush (Viburnum alnifolium) 



same town, both of the last two 



more 



Harris 



Flora by adding numerous species that were either not 
present or had not been reported in 1880, and it gives much 
additional information not included by Robinson. It fur- 
nishes extensive coverage for an area that has lacked ade- 



me 



It also stands as a testimonial to the competence of Stuart 
Harris as a taxonomist, and the sections contributed by 
others serve as a fitting memorial to a highly respected 
botanist whose life was all too short. 



JESUP HERBARIUM 

DEPARTMENT OF BIOLOGICAL SCIENCES 

DARTMOUTH COLLEGE, HANOVER, N.H. 03755 



CAREX GEYERI: REVISITED 



Paul E. Rothrock 



In this journal over 35 years ago, R. T. Clausen and 

H. A. Wahl (1939) first reported the occurrence of Car ex 

geyeri Boott (a typically Rocky Mountain sedge ranging 

from British Columbia and Alberta to Colorado, Utah, and 

northern California) in the Eastern United States. This 
site: 

By limestone outcroppings, in dry deciduous 
woods on bluffs and slopes on west side of Spring 
Creek near west boundary of grounds of State 
Penitentiary, Rock, north of Lemont, Centre Co. 
[Pennsylvania] 

apparently still represents the only known locality for this 
species east of Colorado. 

With the assistance of Dr. Wahl, I was able to relocate 
Carex geyeri in May, 1975. The area has remained undis- 
turbed and, except for sapling growth, mostly unchanged. 
Clumps of C. geyeri were observed in 3 distinct areas 
along a quarter mile stretch of the bluff and have been 
seen at various times in similar habitats slightly down- 
stream (Wahl, personal communication). These clumps 
showed a full range of vigor : fully half were setting fruit, 
others had aborted fruit (perhaps due to lack of proper 
pollination?), while several clumps were only persisting 
vegetatively. 

The bicentric distribution of this sedge may have arisen 
either through introduction by man in recent years or by 
some more natural means. Hermann (1970) favors the 
first hypothesis, but Clausen and Wahl made definite note 
of the natural undisturbed aspect of the habitat. Further- 
more, it should be noted that the nearest major travel 
routes, i.e., railroads, highways, or waterways, are at least 
several miles away thereby making introduction by man 
unlikely. Since other East-West disjuncts are known (see 



538 



1975] Carex Gcyeri — Rothrock 539 



Wood, 1970, for a full discussion) and considering the 
relatively sizable area these plants cover, the localized, dis- 
tinctive nature of the habitat, and the period of time over 
which C. geijeri has been known for this site (first dis- 
covered in 1932), I believe that the occurrence of this sedge 
in central Pennsylvania represents a relict disjunct of 

nhvtoe-eoo-ranhical importance. 



LITERATURE CITED 

Clausen, R. T., & H. A. Wahl. 1939. Plants of central Pennsyl- 
vania. Rhodora 41: 28-34. 

Hermann, F. J. 1970. Manual of the Carices of the Rocky Moun- 
tains and Colorado Basin. Agriculture Handbook No. 374. 
U. S. D. A. Washing-ton, I). C. 397 pp. 

Wood, C. E., Jr. 1970. Some floristic relationships between the 
Southern Appalachians and Western North America. Pp. 331-404 
in: P. C. Holt Ed., The distributional history of the biota of 
the South Appalachians. Part II: Flora. Va. Polytech. Inst. & 
State Univ., Blacksburg, Va. 



202 BUCKHOUT LAB 
PENNSYLVANIA STATE UNIVERSITY 

UNIVERSITY PARK, PENNA. 16802 



INTERNATIONAL REGISTER OF COMPUTER 

PROJECTS IN SYSTEMAT1CS 



Sponsored By 
the international association for plant taxonomy 

and the society of systematic zoology 



Call For Information On Projects, Programs 

And Data Files 



The above two international associations are the prime 
sponsors of an International Register of Computer Projects 
In Systematics. For the purpose of the Register, system- 
atica includes taxonomy, biosystematics, evolution, and 
biogeography of all biological taxa. The Register also 
welcomes information about nonbiological data files of use 
to systematics (e.g., the long range weather data tapes of 
the U.S. Weather Bureau). For the present, our project 
is a Register, which hopefully can direct people to the 
source of information desired. Depending on demand, it 
could be extended into a repository and clearing house for 
computerized files of systematic value. 

As in the first such Register (see Taxon 19:63-76 [1970] ) 
we welcome systematic information on computerized data 
files about living organisms, preserved organisms, experi- 
mental data, literature files, etc. We also welcome informa- 
tion on well-written and documented computer program 
packages (other than basic statistics) that are of value 
for systematic research and/or teaching. 

If you or a colleague use computers in systematics (or 
definitely plan to), please write to the Chairman of the 
Register, and request as many copies of the Register 
Questionnaire as you have separate projects or program 
packages. You will be helping systematics in general by 
avoiding duplication of effort and by contributing to our 
attempts to minimize the incompatibility of computerized 
systematic data, or programs generated on different proj- 
ects. You will be helping yourself because not only might 



540 



1975] Computer Projects — Crovello 541 



you 



discover that someone else has already written the 



com 



programs you have created may 



others, thus enhancing their value. 

The Register will be computerized and available for 
customized search requests by September 1976. As demand 
warrants it, published summaries will also appear. This 
Register will be compatible with a similar Register for all 
of biology that Crovello is organizing for the American 
Institute of Biological Sciences. 



mati 



Q 



THEODORE J. CROVELLO, CHAIRMAN 
INTERNATIONAL REGISTER 
DEPARTMENT OF BIOLOGY 
UNIVERSITY OF NOTRE DAME 
NOTRE DAME, INDIANA 46556 U.S.A 



Volume 77, No. 812, including pages Ul-552, ivas issued Dec. 31, 1975 




JOURNAL OF THE 

NEW ENGLAND BOTANICAL CLUB 



Conducted and published for the Club, by 
ALFRED LINN BOGLE, Editor-in-Chief 

ROLLA MILTON TRYON 
STEPHEN ALAN SPONGBERG 
GERALD JOSEPH GASTONY 
RICHARD EDWIN WEAVER 



Associate Editors 



VOLUME 77 



1975 



Wbt 5fcfa Snglanit liotaittral Club, 3nr 



L. 



Botanical Museum, Oxford Street, Cambridge, Mass. 02138 



INDEX TO VOLUME 77 

Entries include: authors; title words such as geographic area; and 
new records, systematic revisions, or ecological studies by taxa. Lists 
or tables of taxa and maps are also classified spearately under "Lists" 
and "Maps". Note that only one page number per article is given 
for each taxon entered. New scientific names and combinations are 
printed in bold face type. 



Alsophila Brooksii X Nephelea 
portoricensis 451, A. dryopter- 



I5ogle, A. Linn, Editor's Note to 
Contributors 163-164 



oides X N. portoricensis 442, Butomus umbellatus 160 



A. hotteana X Nephelea sp. 

452 

Amorpha, revision of North 
American 337, Key to Species 

340, californica 345, californica 
var. californica, var. napensis 
348, apiculata 350, herbacea 
352, key to varieties of herba- 
cea 357, var. floridana 358, A. 
crenulata 360, georgiana 362, 
key to varieties of A. georgi- 
ana, var. georgiana, var. con- 



Cantino, Philip D., Boerhavia 
spicata (Nyctaginaceae), The 
North American Species, In 
Northwestern Argentina, 423- 

426 
Cape Cod Algae, map 77, Sea- 
sonal occurrence and reproduc- 
tion, figures 78, 79, tables 86- 
96, Distribution, tables 86-102, 
Longevity, table 97-99, Annuals 
and Perennials, tables 100- 103 



fusa 366, A. eanescens 367, nana Carex geyeri 537 

373, glabra 377, paniculata 384, Ceramium fastigiatum, 28, gra- 

nitens 386, laevigata 388, roe- cillimum 27 

meriana 391 ouachitensis sp. Chimaphila maculata, in New 



nov, 394, fruticosa 397 
Antithamnion antillarum 26 
Ascophyllum nodosum 16 
Asterocytis ramosa 22 
Ayensu, Edward S., Report on 

Endangered and Threatened 

Plant Species of the United 

States 355 



Parker, William T., (see God- 

fread, C.) 
Baskin, Jerry M. and Carol C, 
Geographical Distribution of 
the Cedar Glade Endemic Viola 
Egglestonii 427-429 



Hampshire 436 

Chondria 31 

Chordaria flagelliformis 13 

Chromosome Counts of Composi- 
tae From The United States, 
Mexico and Guatemala 171-195 

Cladophora dalmatica, laetevirens, 

6, socialis 7 
Coleman, Douglas C. and Arthur 
C. Mathieson, Investigations of 
New England Marine Algae 
VII: Seasonal Occurrence and 
Reproduction of Marine Algae 
Near Cape Cod, Massachusetts, 
76-104 



Ratrachospermum, Observations Colpomenia sinuosa 13 



in 



Southeastern 



Streams 467 
Boerhavia spicata 423 



Wisconsin Compositae, Chromosome Counts 

of figure 174-175, Table 182- 
191 



545 



546 



Rhodora 



[Vol. 77 



Computer projects in systematics, Erythrotrichia carnea 23 



International register of 539 



Feldmannia irregularis 8 



Conant, David S., Hybrids in Ferns and Fern Allies, A County 



American Cyatheaceae 441-455 
Crotalaria sagittalis, Facultative 

Dwarfism, Notes On the Le- 

guminosae II. 141 
Crouania attenuata 28 



Checklist of in Kansas, Ne- 



braska 



» 



South Dakota and 



North Dakota 478 
Fosliella farinosa, 25; lejolisii, 
25, 26 



Crovello, Theodore J., Inter- Fralick, Richard A. (See Riggs, 



national Register of Computer 



S,A.) 



Projects in Systematics 539-40 Fucus vesiculosus 16 



Crow, Garrett E., Book Review: 
Winter Keys to Woody Plants 
of Maine 438-439 

Cyatheaceae, Hybrids in Ameri- 
can 441 

Cyathea arborea 448, irregularis, 
jamaicensis 453 



Dasya rigidula 30 

Davidse, Gerrit (See Spellman, 

D.L.) 
De Plantis Toxicariis E. Mundo 

Novo Tropicale Commenta- 

tiones XII, Notes on Biody- 

namic Piperaceous Plants 165- 

170 
De Wolf, Gordon P. Jr., The 

Flora of Concord: A Review 

334 

Dictyota sp. 11 
Diplochaete solitaria 4 
Dwyer, John D. (See Spellman, 
D.L.) 



Geranium sibiricum 162 
Giffordia conifera 9; G. littoralis 

10; G. mitchelliae 9; G. san- 

driana 10 
Godfread, Carolyn, Butomaceae: 

A New Family Record for 

North Dakota, 160-161 
Griffithsia radicans 29 



Handlos, Wayne L., The Taxo- 
nomy of Tripogandra (Comme- 
linaceae) 213-333 

Harriman, Neil A. , Geranium 

sibiricum L. (Geraniaceae) in 
Wisconsin 162 

Herposiphonia secunda, 32; te- 

nella 32 
Heterosiphonia wurdemanni 30 
Hypoglossum tenuifolium 31 

Index Holmensis Project, Appeal 

for Support 336 
Instructions to Contributors 551 
Isthmoplea sphaerophora 15 

Jania adherens 24; capillacea, 25 



Ectocarpus elachistaeformis 8 

Elachistea lubrica 12 

Endangered and Threatened Spe- 
cies of the United States, A Keil, David J., Pectis humifusa 

New to the Flora of the United 
States 145-146 



Report on 335 
Enteromorpha sp. 4 
Ericaceace, Bog, Apparent Eco- Keil, David J. and Tod F. Stuessy, 



typic Differences In The Water 
Relations Of Some Northern 53 
Erythrocladia recondita; subin- 
tegra 23 



Chromosome Counts of Com- 
positae from the United States, 
Mexico, and Guatemala 171- 
195 



1975] 



Index to Volume 77 



547 



Kent Island, A Vegetational Sur- 
vey of the Vascular Plants of 
196 

Kiger, Robert W., Papaver in 
North America North of Mex- 
ico 410-422 

Krai, Robert, Rudbeckia auricu- 
lata (Perdue) Krai, A Species 
Distinct From R. fulgida Ait. 
44-52 



Lactuca muralis, in Maine 434; 

in British Columbia 435 
Leguminosae, Notes on the 141 
Lichens, Maritime and Marine 147 



LISTS (TABLES) 

Batrachospermum, localities, 
abbreviations and sampling 
dates 468-469, range in envi- 
ronmental conditions for 472, 
range in selected environ- 
mental conditions for Scup- 
pernong Ck. and Tichigan 
Ck. 472 

Belize Monocotyledoneae 113- 
139 

Botanical Collectors in Belize 
110-112 

Cape Cod Marine Algae Dis- 
tribution, Longevity, Domi- 
nant Annuals and Perennials, 
Seasonal Occurrence, Repro- 
ductive Periodicities 86-103 
Chromosome Counts of Com- 
positae from the United 
States, Mexico, and Guate- 
mala 182-191 

Frequency of Kent Island 

Shrubs and Herbs 201 
Herbaceous and Woody Species 

Cover, Kent Island 207-208 
Marine Algae from Nahant, 

Massachusetts 152-155. 430- 



Marine Lichens from Nahant, 
Massachusetts 147-148 

Seaweed, composition at Dover 
Point, New Hampshire 522- 
525, monthly occurrence and 
longevity of 527-529, num- 
bers of taxa from Dover 

Point 530 
Soil analysis Data for Kent 

Island Transects 204 
Zostera marina, Characteristics 

of eight stations within the 

Great Bay Estuary 457 

Lophosiphonia cristata, 33 



Magnolias, The Deciduous of 
West Florida 64, acuminata 70, 
Ashei 66, pyramidata 71, tripe- 

tala, 72 



MAPS 

Amorpha canescens, Distribu- 
tion of 371, Amorpha spp., 
in Southeastern U.S. 395, in 
Southwestern U.S. and Mex- 
ico 349, fruticosa, in U.S. 
404, glabra, schwerinii and 
paniculata, in U.S. 380, nana, 
in U.S. 376, nitens, laevigata, 
roemeriana, ouachitensis in 

U.S. 389 
Belize, Administrative/Political 

Districts 106 

Kansas, Nebraska, South Da- 
kota and North Dakota, 
County maps 481 

Marine Algae Stations on Cape 



Cod 



( / 



433 



New Hampshire Coast and the 
Great Bay Estuary System, 
map 513, Dover Point and 
surrounding areas 514 

Vegetation Map, Kent Island, 
New Brunswick 198 






548 



Rhodora 



[Vol. 77 



Viola egglestonii, geographical Percursaria percursa 5 



distribution 428 



Marchand, Peter J., Apparent 
Ecotypic Differences in the 
Water Relations of Some 
Northern Bog Ericaceae 53-63 



Petalonia fascia 14 

Petrik-Ott, Aleta Jo, A County 
Checklist of the Ferns and 
Fern Allies of Kansas, Ne- 



braska, South Dakota 
North Dakota 478-511 



and 



Marine Algae of Nahant, Massa- Phragmites communis 159 

chusetts 149-158 Piperaceae (see Schultes, R. E.) 

Mathieson, Arthur C. (see Cole- Piper Allenii, auritum, Bartlin- 



man 



D.C.), (see Reynolds, 



gianum 166; dactylostigmum, 
P. erythroxyloides sp. nov. 167; 
hispidum, P. cf. interitum 169; 
P. sp. 170 
of the Kent Island Group, Plant Collecting in Belize, a 



Norman B.) 
McCain, John W., A Vegetational 
Survey of the Vascular Plants 



Grand Manan, New Brunswick 

" 196-209 

Miller, Ronald F., The Deciduous 



Historical Introduction 105 
Polysiphonia delicatula, lanosa, 
34; poko 35 



Magnolias of West Florida, Poole, James P., Book Review: 



The Flora of Essex County, 
Massachusetts 533-536 
Poole, James P., Chimaphila 
maculata (L.) Pursh in New 
Hampshire 436-437 
Nahant Algae 149, recent addi- Poole, James P., Lactuca muralis 



64-75 
Monocotyledonae of Belize 105; 

list of, 113-139 
Monostroma pulchrum 5 



tions to 430 



(L.) Gaertn. In Maine 434 



Nephelea balanocarpa X Nephe- Pothomorphe umbellata 170 
lea woodwardioides 452 Pseudotetraspora marina 3 

Notes On the Leguminosae II. Punctaria latifolia, plantaginea 



Facultative Dwarfism In Cro- 
talaria sagittalis 141-144 



14 
Pylaiella littoralis 10 



Padina sp. 12 

Papaver, in North America north 
of Mexico 410, Papaver sp. 411, 
key to species 412, somniferum 



Reviews: Flora of Concord 334, 
Flora of Essex County 533- 
536, Winter Keys to Woody 
Plants of Maine 438-439 



413, rhoeas, dubium 414, cali- Reynolds, Norman B. and Arthur 



fornicum, hybridum 415, orien- 
tale 416, alpinum, alboroseum, 
walpolei 417, mcconnellii 418, 
lapponicum 419, nudicaule 420 



C. Mathieson, Seasonal Occur- 

and Ecology of Marine 



rence 



Algae in a New Hampshire 
Tidal Rapid 512-532 



Pectis humifusa, 
U.S. 145 



record for Riggs, Stanley A. and Richard 

A. Fralick, Zostera marina L., 



Peperomia emarginella, glabella 
var. melanostigma 165; serpens 
166 



Its Growth and Distribution in 
The Great Bay Estuary, New 
Hampshire 456-466 



1975] 



Index to Volume 77 



549 



Rothrock, Paul E., Carex geyeri: Thalia geniculata, L. f. rheu- 



Revisited 537-538 
Rudbeckia auriculata, a Species 
Distinct From R. fulgida 44, 



moides Shuey, forma nova, A 
Red-Petioled Form From Cen- 
tral Florida 210-212 



stat. nov. 49; R. fulgida var. Tralau, Hans, Appeal for Sup- 



fulgida, R. fulgida var. um- 
brosa, Plate 45, Habit sketches 
46 



Sargassum, Conspectus of Taxa 
in the western Sargasso Sea 
(key) 17, fluitans 18, hystrix 
19, natans 19, "A", 18, 20; "B", 
18, 20; "C", 18, 21; "D", 17, 21 
Saxifraga Aizoon, S. cernua 41 
Saxifrages on Mount Washing- 



ton 41 

Schultes, Richard Evans, De 
Plantis Toxicariis E Mundo 
Novo Tropicale Commenta- 
tiones XII. Notes on Biody- 
namic Piperaceous Plants 165- 
170 

Scytosiphon lomentaria 15 

Shuey, Allen G., A Red-petioled 
Form of Thalia geniculata 
L. from Central Florida 210- 
212 

Spellman, David L., John D. 
Dwyer and Gerrit Davidse., A 
List of the Monocotyledoneae 
of Belize Including a Historical 
Introduction to Plant Collect- 
ing in Belize 105-140 

Spermothamnion investiens 29 

Sphacelaria fucigera 11 

Spongomorpha arcta 7 



port for the Index Holmensis 
Project 336 



Tripogandra 



(Commelinaceae) 



the Taxonomy of 213-333; 
Historical Review 214; Morph- 
ology 218; Cytology, Breeding 
Systems and Pollinators 231 ; 
Measurements 234; Systematic 
Account 235; Key to the Spe- 
cies of 237; T. amplexans sp. 
nov. 242, amplexicaulis 245, an- 
gustifolia 249, brasiliensis, sp. 
nov. 253, disegra 254, diuretica 
comb. nov. 259, encolea 262, 



glandulosa 



265, 



grandiflora 



268, guerrerensis 272, kru- 
seana 275, montana 276, multi- 
flora, neglecta sp. nov. 287, 
palmeri 288, purpurascens 291, 
purpurascens subsp. purpuras- 
cens 293, purpurascens subsp. 
australis 297, saxicola 298, ser- 
rulata 301, warmingiana 311 

Vegetational Survey of The Vas- 
cular Plants of the Kent Island 
Group, Grand Manan, New 
Brunswick 196-209 

Viola egglestonii, geographical 
distribution of 427 

Watson, Thomas Jr. (see Withe r- 
spoon, John T.) 



Stalter, Richard, Phragmites Webber, E. E., Maritime and 



communis in South Carolina 
159 



Marine Lichens from Nahant 
147-148 



Statement of Ownership, inside Webber, E. E., Phycological Stud- 



back cover, December, 1975 
Steele, Frederic L., Saxifrages on 

Mount Washington 41-43 
Stuessy, Tod F. (see Keil, David 

J.) 



ies from the Marine Science 
Institute, Nahant, Massachu- 
setts I: Introduction And Pre- 
liminary Tabulation of Species 

at Nahant 149-158 



550 



Rhodora 



[Vol. 77 



Webber, E. E., Recent Additions Woelkerling, William J., On the 



to the Marine Algal Flora of 

Nahant, Massachusetts 430-433 

Wilbur, Robert L., A Revision of 

The North American Genus 



Epibiotic and Pelagic Chloro- 
phyceae, Phaeophyceae, and 
Rhodophyceae of the Western 
Sargasso Sea 1-40 



Amorpha (Leguminosae-Psora- Wrangelia argus 30 
leae) 337-409 
Windier, Donald R., Notes on the Xylem Water Potential of Some 



Leguminosae II: Facultative 
Dwarfism in Crotalaria sagit- 
talis L. 141-144 



Bog Ericaceae Figure 57, Diur- 
nal Trend of Water Potential, 

Figure 59 



Witherspoon, 



John 



T. 



and 



Thomas Watson Jr., Lactuca Zostera marina, growth and dis- 



muralis in British Columbia 
435 
Woelkerling, William J., Obser- 
vations On Batrachospermum 
(Rhodophyta) in Southeastern 
Wisconsin Streams 467-477 



tribution in New Hampshire 
456, hydrographic factors 459, 
seasonal growth 460, standing 
crop 461, horizontal transplants 
462 



Missouri Botanical Garden Library 







1753 00343 1308 



1975] 



551 



INSTRUCTIONS FOR CONTRIBUTORS TO RHODORA 
Manuscripts should be submitted in duplicate and should 
be double-spaced or preferably triple-spaced (not on cor- 
rasable bond), and a list of legends for figures and maps 
provided on a separate page. Footnotes should be used 
sparingly, as they are usually not necessary. Do not indi- 
cate the style of type through the use of capitals or under- 
scoring, particularly in the citations of specimens, except 
that the names of species and genera may be underlined to 
indicate italics in discussions. Specimen citations should 



common 



em 



distribution. Sys 

be prepared in the format of "The Systematics and Ecology 
of Poison-Ivy and the Poison-Oaks," W. T. Gillis, Rhodora 
73: 161-237, 370-443. 1971, particularly with reference to 
the indentation of keys and synonyms. Papers of a floristic 
nature should follow, as far as possible, the format of 
"Contribution to the Fungus Flora of Northeastern North 
America. V.," H. E. Bigelow & M. E. Barr, Rhodora 71: 177- 
203. 1969. For I 
of standard journal abbreviations is given by L. Schwarten 



recomm 



W 



1958 



RHODORA December, 1975 Vol 77. No. 812 



CONTENTS 



Hybrids in American Cyatheaceae. 

David S. Conant 



441 



Zostera marina L.. Tts Growth and Distribution in the Great Bay 

Estuary, New Hampshire. 

Stanley A. Riggs f Jr., and Richard A. Fralick 456 

Observations on Batrachospermum (Rhodophyta) in Southeastern 
Wisconsin Streams. 

William J. Woelkerling 467 

A County Checklist of the Ferns and Fern Allies of Kansas, 
Nebraska, South Dakota, and North Dakota. 
Aleta Jo Petrik-Ott 478 

- 

Seasonal Occurrence and Ecology of Marine Algae in a New 

Hampshire Tidal Rapid. 

Norman B. Reynolds and Arthur C. Mathiemm 512 

Book Review: The Flora of Essex County, Massachusetts. 

James P. Poole 534 



Carex geyeri: Revisited. 

Paul E. Rothrock , 



538 



International Register of Computer Projects in Systematica. 

Theodore J. Crovelio 540 

Index to Volume 77 545 

Instructions to Contributors 551 

COVER III. Statement of Ownership. 



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2 DATE OF FILING 

S ept. 23,1975 



4 LOCATION OF KNOWN OFFICE OF PUBLICATION (Strret. city, county, state and ZIP code) (Not printer,, 



: 



3A. ANMLIAL SUBSCRIPTION PRICE 
$20.00 



New E ngland Botani ca l Club, Inc., Bot anical Museum, Oxford Street, Cambridge, Mass . 02138 

5 LOCATION OF THE HE ADQUARTERS OR GENE R AL BUSINESSOF F ICES OF THE P U^^rRSiNoTpTin t e rs , " ~~~ ~ 

[. Npw FnglanH Rnfanira i r.1 nh, Tnn . f Botanical Museum , Oxford Street # Cambridge, Mass . 02138 

NAMES AND ADDRESSES OF PUBLISHER, EDITOR, AND MANAGING EDITOR 



6 



PUBLISHER (Name and address. 



New England Botanical Club, Inc ., Botanical Museum, Oxford Street , Cambridge, Mass. 02138 



EDITOR (Name and address, 

Alfred Linn Bogle, Dept. Botany , University of New Hampshire , Durham, N.H. 03824 



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erman R. Sweet, Botanical Museum, Oxford Street, Cambridge , Mass . 02138 




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